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OF THE 


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ONS AND PROCEEDINGS 
OVAL SOCIETY of SOUTH AUSTRALIA 


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TRANSACTIONS AND PROCEEDINGS 


OF THE 


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[With Forty-two PLATES, AND SIXTY-sIx 
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Vi. 


Ropul Society of South Australia 


(INCORPORATED). 


Patron: . 
HIS EXCELLENCY LIEUT.-COL. SIR G. T. M. BRIDGES, 
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- PAGE 


Ossporn, Pror. T. G. B.: A Note on the Pathological 
Morphology of Cintractia seus, ee) MeAlp. 
Piate t..2-%:. 

Davin, Pror. Sir T. Epcrworrn: Gecumecnes ie Bewains a 
Small Crustacea in the Proterozoic (?) or Lower Cam- 
brian(?) Rocks of Reynella, near Adelaide. Plate ii. 

Asusy, Epwin: Notes on Australian Polyplacophora, with 
Descriptions of Three New RuEeL and Two New Varie- 
ties. Plate iii. 


CuHILton, Pror. CHARLES: ms ek Teena. en ‘Contral ae: 
tralia belonging to the Phreatoicidae 

Cuitton, Pror. Cuartes: The Flora and Fauna He Ng 
Archipelago and the Investigator Group, No. !—The 
Amphipoda and Isopoda 

Jonrts, Pror. F. Woop: The External Cae ae Patch 
Kmbryos of Marsupials, No. 3—Isoodon barrowensis 

Faget hae Pror. WatteR: <A _ Geological Traverse of the 
Flinders Range from the Parachilna Gorge to the Lake 
Frome Plains. Plate iv. 

PuLLEINE, Dr. R. H.: Two New Species of ie fon Sous 
Australia. Plate v. 

CLELAND, Pror. J. Burton: The ecnstiees of Mente aioe Age 

JONES, Pro. F. Woop: The External Characters of Pouch 
Embryos of pope pialss No. 4—Pseudochirops dahli. 
Plate vi. ... ee oe ra pe * 

Mawson, Pror. Sig eons “The Jectiary Brown-coal 
Bearing Beds of Moorlands 

Rogers, Dr. R. S.: Contributions ie ae Orchidology of 
Australia and New Zealand 


TraLe, Dr. E. O.: The Physiography of ‘the Miasdays Vallone 
Mount Lofty Ranges 


~Osporn, Pror. T. G. B., and Ctomine ee dune New 
Records of Fungi for South Australia, Part II., together 
with a Description of a New Species of Puccinia. 
Plate vii. igs oe ae neg ie oe 

Jonrs, Pror. F. aoa The Flora and Fauna of Nuyt’s 
Archipelago and the Investigator aeote: No. 2—The 
Monodelphian Mammals ; ee ee co 

Ossorn, Pror. T. G. B.: Flora sna Fauna of Nuyt’s 
Archipelago, No. 3—A Sketch of the neat of Franklin 
Islands. Plates viii. to xi. 

Berry, Purare A.: An Investigation of ae iieseatial Oil 
from Eucalyptus cneorifolia, DC. (the ‘‘Narrow Leaf 
Mallee” of Kangaroo Island) Lee 

Trees, O. W.: On the Arrangement of ihe Gorigdons oF 
Voluntary Muscle Fibres in Double Spirals. Plate xii. 


Turner, Dr. A. Jerreris: Australian Fe oben of the 
Group Geometrites 

Lea, Anruur M.: The Flora and Fauna of ‘Nuyt’: s Archi- 
’pelago and the Inv pptaee tor oe No. ry es ia 
Plate xiii. 


85 


119 
131 
148 


160 


166 


181 


viii. 


CONTENTS (ConrinueEp). 

a ; Page. 

PuLLeInE, Dr. Rospert: Cylindro-conical and Cornute Stones 
from the Darling River and Cooper Creek. Plate xiv. 304 

Kuston, Atsert H.: Australian Coleoptera, Part III. ... 309 

Tires, O. W.: Researches on the Insect Metamorphosis, 
Part I.—On the Structure and Post-embryonic Develop- 
ment of a Chalcid Wasp, Nasonia. Part Il.—On the 
Physiology and Interpretation of the Insect Meta- 


morphosis. Plates xv. to xxx. be: Mrs % sx, -Aeee 
Noses, E. Dorotny: A Preliminary Note on the Fossil 

Woods from some Australian Brown Coal Deposits... 528 
TrnpaLe, Norman B.: On a New Genus and Species of Aus- 

tralian Lycaeninae. Plate: (Exein. 22. 537 
Apamson, R. S., and Pror. T. G. B. Osporn: On the Beology 

of the Ooldea District. Plates xxxii. to xxxvi. ... 539 
Buack. J. M.: Additions to the Flora of South Auseealiaee 

No. 20. Plate xxxvii. 565 


AsuBy, Epwin: Types of sya of AREA ase Payne 
cophora described by de Blainville, Lamarck, de Reoche- 
brune, and Others, now in the Museum a Beto 


Naturelle, in Paris... 572 
Istnc, E. H.: Ecological Notes on : Soe eect a ie Plants, 
Part: Plates xxxviii. to -xlii; ue 583 
MISCELLANEA ... ss Be oe — ne ee 
ABSTRACT OF Buacsiiiees ae iy oe at ieack sas ge 
PRESIDENTIAL ADDRESS ae ee aa a noe adi) Oe 
ANNUAL REPORT PON koe =e ae ae ye «a; GOenD 
BaLANCE-SHEETS ay 5 mt, vty a s: it 6S 
Donations To LIBRARY ae ea ke r ces ... 684 
List or MremBers ey a a nek oi > 3) 664 
APPENDICES : — 
Field Naturalists’ Section: Annual Report, etc. — 667 
Thirty-third Annual Report of the Native Fauna and 
Flora Protection Committee... 669 


INDEX -<.. a ye Nee fe bah ee a Pe: bar taae e 


INDEX TO THE TRANSACTIONS. 


The following works can be had, at the prices mentioned, by 
applying to the Hon. Secretary of the Royal Society, Royal Society 
Rooms, North Terrace, Adelaide :— 


INDEX TO THE TRANSACTIONS, PROCEEDINGS, AND REPORTS OF 
THE Roya Society or Sovutu AUSTRALIA, Vols. I. to 
XXIV., 1877-1900. 


InDEX 10 tHE TRANSACTIONS, PRockEDINGS, Rwports, AND 
Memoirs OF THE Roya Socrery or SovutH AUSTRALIA, 
Vols. XXV. to XLIV., 1901-1920. 

Price: Five Shillings each volume; to Fellows and 
Members, hialf-price. 


THE 


Transactions 
OF 


The Royal Society of South Australia 


(Incorporated.) 
‘Vol. XLVI. 


A NOTE ON THE PATHOLOGICAL MORPHOLOGY OF 
CINTRACTIA SPINIFICIS (LUDW.) MCALP. 


By T. G. B. Ossorn, D.Sc., 
Professor of Botany in the University of Adelaide. 


[Read November 10, 1921.] 
Puare 2. 


The fungus now known as Cintractia spinificis was 
described, in 1893, by Ludwig from material collected by Mr. 
J.G.O. Tepper, near Port Adelaide. In the original descrip- 
tion the fungus, which was placed in the genus Ustilago, was 
stated to occur on the female inflorescences, destroying the 
ovaries. 

In his monograph on ‘‘The Smuts of Australia’’ (1910) 
McAlpine redescribed the fungus, transferred it to the genus 
Cintractia, and gave an account of the method of spore forma- 
tion and germination. 

The purpose of this note is twofold—first, to place on 
record the presence of the fungus in the male inflorescences; . 
and, second, to describe certain modifications of the host, 
occurring in both male and female inflorescences, due to the 
presence of the parasite. 

Cintractia smnificis was first noted on the male inflores- 
cences of Spinifex hirsutus, in February, 1918, at Wright 
Island, Encounter Bay. The season was then far advanced, 
and almost all of the spores were shed; there was, however, 
sufficient evidence to determine the fungus provisionally. In 
subsequent seasons—January, 1919 and 1920—it has been 
found in abundance at Victor Harbour, occurring on the male 
as frequently as on the female inflorescences. It has also been 
found at Grange, not far from the type locality in which 


2 


Tepper first collected it on the female. The smut is less con- 
spicuous on the male inflorescences, nor is so large a spore 
mass formed, which may account for it being overlooked by 
previous collectors. 


PATHOLOGICAL CHANGES IN THE Host. 


The inflorescences of the ‘‘spiny rolling grass’’ are common 
and conspicuous objects along the coastal dunes of South 
Australia. The salient features of the normal inflorescences 
will first be described, then the pathological changes noted. 

The normal male inflorescence is a roughly spherical 
head, about 15 cms. in diameter, borne on a stout upright 
stem. It consists of an aggregation of stiff secondary axes, 
each arising in the axil of a bract, upon which the spikelets 
are borne. Below the terminal head there is usually one, 
sometimes two, smaller lateral clusters of secondary axes. 
Each secondary axis is a stout structure about 7 or 8 cms. 
long, bearing a group of 10 to 20 irregularly spirally arranged 
spikelets distributed over the middle third of its length. The 
upper and lower portions of the axis bear no spikelets, the 
upper part terminating in a stiff tapering spine. The spikelet 
is composed of two sterile glumes and two flowering glumes, 
or three sterile and one flowering glume. The flower consists 
of glume, pale, two broad lodicules, and three stamens. The 
axis of the flower ends abruptly with the stamens: no ovary 
rudiment has been seen in the flowers examined. 

The smutted male inflorescences are much slenderer and 
more diffuse (pl. 1., fig. 1). The main differences are: — 

(a) Greater elongation of the internodes of the upright 

stem. In the smutted specimens the average 
length of internode between the terminal head and 
the small lateral cluster, next below it, was 
9°4 cms. as against 6°7 cms. in healthy specimens. 
(6) Reduction in the number of secondary axes bearing 
spikelets in the inflorescence; an average of 16 per 
head in the smutted specimens as against 64 in the 
healthy specimens. | 

(c) The closer aggregation of the spikelets and an increase 

in their number per secondary axis. 


The smutted male spikelets consist of the two sterile 
glumes and two florets. Each has the fertile glume, pale, 
and three stamens. No lodicules were seen (text fig. 1), the 
anthers are about normal length, but contain no pollen, and 
the filaments do not elongate. No ovary recognizable as such 
is present, but the axis of the flower elongates above the point 
of stamen insertion, producing an irregular conical mass 


3 


1-7 mm. long. This, in the ripe smut gall, consists of a 
central core of host tissue coated by the spore mass, which is 
bounded externally by the usual white skin seen in Cintractias. 

As is well known, the normal female inflorescence of 
Spinifec hirsutus is a large globular head, consisting of 
radiating spines, 40 cms., or even more, in diameter. This 
is borne, terminally, on a stout erect stem, and below it is 
one, or sometimes two, small lateral groups of secondary axes, 
bearing flowers. The head itself is not more than 4 or 5 cms. 
above the node below, and when ripe is readily detached by 
snapping off the axis at an absciss region immediately above 
the node. It then blows away, distributing the fruits as it 
breaks up. The head is a complicated system of secondary 
‘axes, arising in the axils of chaffy bracts, about 8 cms. long. 
The majority of these axes are long tapering spines, averaging 
"17 cms. (13°5-20 cms.), which are sterile. They are borne 
in groups of 6-12 or more, each group representing a branch 
system with exceedingly short internodes. These spines form 
the spring-like “‘legs’’ of the tumble weed, and are a most 
characteristic feature of the plant. In each group of sterile 
spines are a few (1-4) shorter and stouter spines, 10-12 cms. 
long (text fig. 2). These are the fertile secondary axes, each 
of which has a single spikelet at its extreme base. The spikelet 
consists of three sterile glumes (or two sterile and one abortive 
male flower) and one fertile glume. The flower has glume, 
pale, three stamens with minute anthers borne on filaments 
as long as the ripe grain, and an ovary. 


The diseased female inflorescence is strikingly different 
from the normal (pl. i., fig. 2). The main differences are: — 


(a) Elongation of the internode below the terminal head. 


(6) Complete absence of the long sterile spines which are 
so obvious in the normal inflorescence. A few 
sterile spines may be present, but these are shorter 
than the fertile spines, of which the head is largely 
built up. 

(c) The spikelets are borne 1°5-4 cms. above the base of 
the fertile secondary axes, which are half as long 
again as normal, z.e., up to 15 cms. 

The smutted female spikelet consists of two sterile glumes 
and two fertile. Both the florets are much modified by the 
fungus (text fig. 4), but the modifications are the same in each 
flower, 1.e., the lower floret, normally an abortive male, 
behaves like a female. The florets have glume and pale, both 
longer than usual, the latter being often involved in the smut 
gall. No stamens have been recognized, the whole of each 
floral axis above the pale being one elongate, rarely purarcate, 
smutty mass (text fig. 5). 


Fig. 1. Smutted male spikelet with two flowers; the glumes 
are all cut away. The stamens are sterile, the filaments do not 
elongate, but the anthers are not smutted. A central smut-body 
has formed (stippled). x5. 


Fig. 2. Normal female secondary axis, with spikelet at its 
extreme base. About natural size. 


Fig. 3. Smutted female secondary axis, of greater length 
than normal, with the diseased spikelets 2 cms. above the base. 
About natural size. 


Fig. 4. Smutted female spikelet with two flowers, the glumes 
all being cut away. The pales are left, that of the upper flower 
being smutted. Note the elongate smut mass that replaces the 
ovary (stippled). x95. 

Fig. 5. Smutted femalé spikelet as above. The lower sterile 
glume is removed and the lower fertile glume is partly cut away. 
The upper pale is hypertrophied and involved in the smut gall. 
The galled ovary of this flower is bifureate. x5. 


5 


GENERAL. 


Various observations as to the effect of parasitic fungi 
upon the flowers of the host are summarized by von Tubeuf.® 

Owing to the extent of gall formation induced by the 
fungus, it is not possible to say if ovaries are actually developed 
in male flowers of Sponifex hirsutus (ef. the cases cited by 
Tubeuf of Carex praecox with U. caricis, Buchlée dactyloides 
with 7. buchléeana, and Andropogon provineialis with U. 
andropogoms). But in place of the normally abbreviated 
floral axis a more or less extensive smut gall is developed, 
resembling that formed in the female flower, except that in 
the male inflorescences it is usually somewhat smaller. A 
similar prolongation of the axis is seen in the axil of the third 
glume of the female spikelet. This glume, it will be remem- 
bered, is either sterile or subtends a male flower in the healthy 
inflorescence. ‘Bas ; 

In a paper on Tilletia foetens, Barrus() cites observa- 
tions by Edler, Appel, and Miczynski upon wheat affected by 
stinking smut to the effect that the diseased heads are looser | 
than normal and of greater length, though Barrus’ own 
observations showed that the infected heads were’ rather 
shorter. He notes that more grains are found in a smutted 
ear of wheat than in a normal one of equal length, there being 
more ovaries per spikelet in the former case. So in Spinfex 
hirsutus two smut masses form per female spikelet, though 
the healthy flower has but a single ovary. 

The most obvious pathological deformation is the com- 
plete absence of the long sterile axes or spines of the normal 
female inflorescence. Thus the smutted head has not the 
same potentiality for distribution as the healthy one, for it 
cannot roll about in the same way. It was probably this 
feature that led to the earlier recognition of the fungus on 
the female plants. More interesting is the development of 
structures resembling those formed in female flowers upon 
the male, which, though they are not so definite as in the 
case of the smuts, referred to above, yet seem to merit brief 
description. 


DESCRIPTION OF PLATE TI. 

_ Fig. 1. Male inflorescence of Spinifex hirsutus infected with 
Cintractia spinificis, showing the reduced number of spikelet- 
bearing axes and an increase in the number of spikelets above 
the normal. : 

Fig. 2. Female inflorescence of S. hirsutus with C. spinificis, 
showing the looser structure, the absence of long sterile spines, 
and the spikelets some 2 cms. above the base of the spikelet- 
bearing axes, abnormalities due to the presence of the fungus. 


(1) Tubeuf and Smith, ‘‘Diseases of Plants,’’ 1897, pp. 26-29. 
(2) Barrus, M. F., ‘“‘Observations on the Pathological Morph- 
plony of Stinking Smut of Wheat,’’ Phytopathology, vi., pp. 21-28, 
916. 


OCCURRENCE OF REMAINS OF SMALL CRUSTACEA IN THE 
PROTEROZOIC (?) OR LOWER CAMBRIAN (?) ROCKS OF 
REYNELLA, NEAR ADELAIDE. 


_ By Proressor T. Epceworta Davin, K.B.E., C.M.G., 
D.S.0., B.A., F.R.S., Hon. D.Sc. Oxford and Manchester. 


(Read November 10, 1921.] 


Puate IT. 


‘DESCRIPTION OF FOSSILS. 


Through the kind assistance of Professor Walter Howchin, 
F.G.S., I was enabled, over a year ago, to examine some 
good sections of the siliceous limestones underlying the 
Brighton limestone, at Reynella, 17 miles southward of Ade- 
laide. The siliceous limestone, as exposed in several of the — 
small quarries belonging to the South Australian Portland 
Cement Company, and nearest to Reynella, on the left bank 
_ of the Field River, shows curious small ochreous bodies in 
a bluish-grey ground-mass. The limestone is mostly oolitic 
in structure. These yellowish-brown to ochreous bodies are 
seen under the microscope to be distinctly of organic origin, 
and there can be little doubt that they are referable to some 
kinds of minute crustacea. Their Bee appearance is shown 
on fig. 3 of pl. u. 

The larger object shown on the left side of fig. 3, and 
about 2 mm. in length, has all the appearance of being a 
swimming paddle. The object marked (f) is possibly part 
of a spiral gill. The remainder of the objects in fig. 3 are 
probably locomotary appendages. 

Fig. 2 probably represents a small carapace. Fig. 1 is 

the only specimen which shows some bilaterally symmetrical 
organization. At the top are traces of what may be antennae 
or antennules, followed below by two pairs of small processes, 
and below these is a pair of stouter appendages, probably 
claws. The spiral object to the right of the claw (?) may be 
one of the spiral gills. A pair of possible parapodia follow, 
and then two fragments of what may have been a somite, or 
body ring. The remainder of the dark objects seen are quite 
problematical. Similar but less well-preserved objects occur 
in the overlying Brighton limestone. 


7 


GEOLOGICAL Horizon. 

The siliceous limestones of Reynella are about 2,500 ft., 
possibly more, below the base of the Cambrian limestones con- 
taining Archaeocyathinae at Sellick Hill, 35 miles southerly 
from Adelaide. The siliceous limestones of Reynella and the 
Brighton limestones, together with those of Burra (on the 
horizon of the Brighton limestones north of Adelaide) are 
singularly like those of the Nullagine series in Western Aus- 
tralia, and called by the Government Geologist (Mr. A. 
Gibb Maitland) the Carawine limestones. The thick reddish- 
purple slate beds of the Adelaide region immediately above the 
Brighton limestone have a close analogy in the reddish- 
purple slates and shales of the Hamersley Range of the 
Fortescue River area of Western Australia and the reddish- 
purple slate series underlying the Irwin River coal measures 
(Greta coal measures), both occurring in the Nullagine series 
of Western Australia. 

The black shales, at least 1,500 ft. thick, which at Sellick 
Hill underlie the Archacocyathinae limestones, and there con- 
tain small chalcedonic nodules, appear to correspond closely 
with the black shales with chalcedonic nodules at the top of 
the Nullagine series of Western Australia, as seen in the 
hills about 16 miles northerly from Roy Hill Station, on the 
Fortescue River. No fossils have, as yet, been found in the 
Nullagine series, and Mr. A. Gibb Maitland classifies them now 
as Proterozoic. 

With the exception of some doubtful radiolaria, figured 
by Professor Howchin and myself from the horizon of 
these siliceous limestones near Hallett Cove,@® and a 
problematical calcareous fossil found by Professor Howchin 
apparently weathered out of the purple slate beds near Hallett 
Cove, no organic remains have previously been recorded from 
these beds in South Australia: Mr. A. Gibb Maitland, as 
already stated, classes the Nullagine series as Proterozoic, 
while Professor Howchin classes these equivalents (in my 
opinion) of the Nullagine series of Western Australia as 
Lower Cambrian. I would tentatively suggest that all the 
strata from the base of the Archaeocyathinae limestones 
to the basal conglomerates overlying the Archaean (?) 
schistose rocks of Aldgate, in the Adelaide region, be 
given some local name such as ‘‘the Adelaide series,” and, 


(1) Proc. Linn. Soc. N.S. Wales, 1896, pt. 4, pp. 571-583, 


pis. xxxix.-xl. 


8 


for the present, I would suggest that they may be classed, 
provisionally, as Proterozoic(?). It is quite possible that 
more than one series of rocks are included in the suggested 
Adelaide series. 

If the crustacean remains, referred to in this paper, are 
really, as I believe, Proterozoic in age, it would be of quite 
extraordinary interest to secure a complete fossil specimen. 
What appear to be casts of annelid burrows can be discerned 
in the weathered outcrops of those remarkable ‘‘varve’’ rocks, 
the Tapley Hill shales, near Adelaide, which occur several 
hundreds of feet below the Reynella horizon. There is, there- 
fore, convenient to Adelaide, a considerable thickness of strata 
containing traces of obscure organisms, and it is to be hoped 
that patient search by local geological workers will soon be 
rewarded by the discovery of some complete specimens. Such 
a discovery would doubtless Pe of priceless value to the 
palaeontologist. 


DESCRIPTION OF PLATE II. 


Remains of small Crustacea from the Proterozoic (?) or Lower 
Cambrian (?), Adelaide Series, Reynella, near Adelaide. 


Fig. 1. Bilaterally symmetrical organism, probably Crusta- 
cean, showing antellules, claws, spiral gill, parapodia(?), ete. 


Fig. 2. Probably a small carapace. 


Fig. 3. Various small appendages, probably locomotary, but — 
(f) may be portion of a spiral gill. 


NOTES ON AUSTRALIAN POLYPLACOPHORA, WITH 
DESCRIPTIONS OF THREE NEW SPECIES AND 
Two NEW VARIETIES. 


By Epwin Asupy, F.L.S., M.B.O.U. 
[Read April 13, 1922.] 
Puate IIT. 


Genus AcanrHocuitTon (Gray, 1821, em.). 


In our paper of October, 1898 (Trans. Roy. Soc. 8S. Austr.), 
and subsequent papers, Dr. Torr and the writer followed Dr. 
Pilsbry in adopting the name for this genus of 4 canthochites, 
Risso, 1826. 

In Proc. Mal. Soc., vol. xi., pt. i1., pp. 126 and 127, 
June, 1914, Mr. Tom Iredale draws attention to the fact that 
E. Gray’s name of Acanthochitona (Lon. Med. Repos., vol. 
xv., 1821) antedates Risso’s name by five years, and he therein 
also points out that in the name commonly used, Risso’s 
spelling has been amended. Iredale followed this in July, 
1915 (Trans. N. Z’d Inst., vol. xlvii., p. 422, 1914), by 
amending Gray’s name in dropping the terminal ‘‘a’’ and 
writing the genus ‘“‘Acanthochiton (Gray, 1821, em.)”. It 
will also be seen that Dr. Pilsbry, /.c., while adhering to 
Risso’s name, gives “Acanthochiton, Herrmannsen, Indicis 
Generum Malacozoorum Primordia, i., p. 2; Acanthochiton 
of Carpenter and many modern authors.’’ While several 
Australian workers have adopted Gray’s name in place of 
Risso’s, as pointed out by Iredale, /.c., it is regrettable that 
they have not followed Iredale in the amended spelling, 
changing Acanthochitona into Acanthochiton. 

In all my papers, in 1918 and since, I have adopted 
Iredale’s spelling for the following reasons: —(1) The terminal 
“Chiton’’ is in keeping with so many other genera.. (2) The 
two words, ‘‘Acantho’’ and ‘“‘Chiton,” are both masculine, 
and Gray seems to have tacked on to these two Greek words 
a Latin feminine terminal, thereby producing what my friend 
the classical professor terms, a ‘‘mongrel word.’’ It is pos- 
sible that Gray added the ‘‘a’’ to make his new genus agree 
with a certain specific name, a course that can hardly be 
justified ; anyhow, I cannot see how we can do otherwise than 
adopt Iredale’s suggestion and drop the unfortunate terminal. 
This course seems to be the commonsense one, and is probably 
the only correct one, and, as Iredale says, there are plenty 
of precedents. 


10 


ACANTHOCHITON (NOTOPLAX) GABRIELI, n. sp. 


Differs from A. costatus, Ad. and Ang., in having deep, 
broken, longitudinal grooving in the dorsal area, whereas 4. 
costatus has smooth, except for the transverse ribs following 
the lines of growth. 

The pustules in the diagonal ribs of the species under 
description are larger, more rounded, and irregular than in 
costatus, and in addition on some of the valves there are 
evidences of a second coarsely pustulose rib on the posterior 
margin of the median valves, in this respect approaching its 
Western Australian ally, A. sub-viridis, Torr. 

Remarks.—The specimen described above has been kindly 
lent to me for description in this paper by my friend Mr. 
Gatliff; it is from Caloundra, in Queensland, and the type, 
which has not been disarticulated, remains in Mr. Gatliff’s 
collection. I am naming it after my friend Mr. Charles J. 
Gabriel, who has for so long been associated with Mr. Gatliff 
in the excellent conchological papers they have jointly pro- 
duced. While I suspect this form should hold sub-specific 
rank only, it is as much entitled to the higher rank as are 
some of its allies referred to in the discussion following. 


ACANTHOCHITON costTaTus, Ad. and Ang., and its allies. 

(Ad. and Ang., P.Z.S., 1864, p. 194; Angas, l.c., 1867, p. 224.) 

The examination of the preceding species from Caloundra 
and, subsequently, the loan of a specimen from Port Philip, 
Victoria, measuring 74 mm. in length, by Mr. Gabriel, has 
made it necessary to go into the whole question of the respective 
relationships of several nearly-related species. 

My series of A. costatus include three from Sydney, 6, 
12, and 19 mm., respectively; several from Tasmania, up to 
36 mm. in length; and one from South Australia, 21 mm. 
All have minute, slender, girdle spicules, easily detached ; all 
have several ribs composed of coarse pustules, behind the 
mucro, in tail valve, but in the smallest this feature is repre- 
sented only by a single large pustule at the outer edge; all 
probably, in the quite juvenile stage, possess none of these 
posterior ribs. In the larger specimens the original form of 
the dorsal area in the juvenile is not quite clear, but in the 
three from Sydney Harbour, 6, 12, and 19 mm. respectively 
(dry), it certainly commences with a prominent, broad, 
rounded beak, the area rapidly widening with sundry jags at 
each side, giving the pinnatifid character. On reaching a total 
length of the whole shell of 6 to 10 mm., the dorsal area may 
slightly contract or continue in two parallel lines, forming in 
the adult a narrow raised dorsal ridge. On collecting the 


11 


small specimen at the Quarantine Station, Sydney, in 1918, 
I at first thought it a different species owing to its wide 
diverging dorsal area, but at last noted that it was only a 
juvenile character. 

Mr. Gabriel’s small shell from Port Philip, about 74 mm., 
which I understand has been recorded as A. rubrostratus, 
Torr, is similar to the juvenile costatus in having small spicules 
clothing the girdle and in having the broad dorsal area of 
the juvenile, but it differs in not having coarse pustulose 
ribbing behind the mucro and in the pustules of the diagonal 
ribs being small; in these two respects only does it resemble 
rubrostratus and speciosus. I believe that had it attained a 
larger growth it would have been quite similar to typical 
costatus in these respects. It may be that in Victoria a race 
of costatus is living that attains the senile characters at a later 
age than is common to that species. 

Throughout this paper all measurements given are of dry 
specimens. I have two specimens collected by myself in Gulf 
St. Vincent, measuring 8 and 18 mm. long, and I have com- 
pared them with Dr. Torr’s type and co-type of A. rubro- 
stratus, and find them con-specific; the girdle is covered with 
coarse white spicules, similar in size to those clothing the 
girdle of A. speciosus, H. Adams, but appear less irregular in 
their attachments and show between the valves in a much less 
degree; the dorsal area is similar in the two, except that the 
pinnatifid character is more continuous in rwbrostratus ; 
neither have ribs behind the mucro, although one of Dr. 
Torr’s latter shows three waves or undulations corresponding 
with the posterior ribbing of costatus, nor have either the 
coarse pustules of that species in the diagonal ribs. The 
width of-the girdle of rubrostratus is less than that of 
speciosus, and the beak of speciosus is less pronounced, but 
when it is considered that the whole of our specimens of 
rubrostratus are smaller than the smallest of our specimens 
of the other species, is it not possible that the slight differences 
enumerated may be due to the juvenility, and that rwubro- 
stratus is really the young of A. speciosus. 

In conclusion.—As before mentioned, I have been much 
indebted to Dr. Torr for the opportunity of examining speci- 
mens in his collection, but until a much larger series of all 
the species, in all stages of growth, is available, I do not 
like to make a final decision, but I am inclined to think that 
ultimately we shall be able to recognize A. costatus as the 
dominant species, with two sub-species—gabrieli, Ashby, from 
Queensland, and sub-viridis, Torr, from Western Australia— 
A. costatus, sub-species, being found in New South Wales, 
Victoria, Tasmania, and South Australia; A. speciosus, 


12 


H. Adams, as a dominant species with rubrostratus, Torr, 
either as a sub-species or as a synonym, being the name distin- 
guishing the juvenile form. 

It should be mentioned that some of the forms above 
referred to show sub-cutaneous lining and others short longi- 
tudinal rows of shallow holes near the beak on the dorsal area. 
The following reswmé may be helpful : — 

_ Minute, slender, girdle spicules: gabrieli, costatus, sub- 
viridis. 

Coarse, girdle spicules: speciosus, rubrostratus. | 

Broad, pinnatifid, dorsal area in juvenile: gabrielt and 
costatus. 

Narrow, pinnatifid, dorsal area: speciosus, rubrostratus, 
sub-viridis. | 

Small pustules and no ribs behind mucro: speczosus and 
rwbrostratus. 


ACANTHOCHITON MAYI, 0. sp. 


Introduction.—A number of median valves of a rather 
striking Acanthochton were dredged by Mr. W. L. May in 
various parts of Tasmania, in depths varying from 60 fms. to 
100 fms. It is such a distinct species and the valves are so 
well preserved that one seems well justified in describing it 
without waiting for the discovery of the whole shell. I have 
much pleasure in calling it after Mr. W. L. May, the dis- 
coverer, and a gentleman who has done such splendid work in 
conchology. 

Specimen No. 1. Type. 


Median valve.—Colour pale cream, very strongly carin- 
ated, prominently beaked, dorsal area well defined, fairly broad 
but sides almost parallel, 2.e., after attaining half-growth the 
sides of this area do not diverge. The central ridge and the 
two sides of this area continue as smooth ribs the whole length 
of the valve; the space between is cut up by short, deep, 
longitudinal grooves, reminding one of cuneiform characters. 
These deep grooves make the ribs, before referred to, jagged 
at their margins. The lateral area is separated from the 
pleural by a fold surmounted by extra large pustules, which 
are widely spaced. The first three rows next the beak are 
composed of minute pustules which quickly lose themselves in 
the lateral rib of the dorsal area, before referred to; the fourth 
row of pustules is composed of three minute and six large ones 
placed diagonally in the row which is parallel to the dorsal area, 
beyond the ninth pustule. In this row the sculpture is con- 
fluent, forming a broad flat rib for about one-third of the 
total length of the valve. The rest of the valve is decorated 
with five rows and one-half row of elongated, much raised, 


13 


flat pustules, widely spaced, and the rows widely separated 
from one another. All pustules are placed on the diagonal, 
and the rows themselves become more and more diagonal as 
the margin of the shell is approached. Slit 11 modified into a 
deep groove, with raised edges on the upper side of the 
articulamentum. 

Measurements.—-3 mm. longitudinally, 3°75 mm. laterally. 

Habitat.—A number of valves dredged 7 miles east of 
Cape Pillar, North-west Tasmania, in 100 fms.; one valve, 
about the same depth off Schouten Island; one valve, coloured 
red, off Port Arthur, South-east Tasmania, in 60 fms. 


Specimen No. 2. Co-type. 


This median valve was likewise dredged in the same depth 
east of Cape Pillar. While corresponding, in the main, with 
the preceding valve, it differs in that the dorsal area is uni- 
formly covered with short, deep, wedge-shaped, longitudinal 
grooves. Also the fold separating the pleural from the lateral © 
area is very strongly raised, and the pustules in the rows, 
where they pass over this fold, are larger and broader than 
anywhere else, for a width of two pustules. If these pustules 
had not been so widely spaced one would have described the 
valve as possessing a wide diagonal rib composed of two rows 
of pustules. In method of sculpture and shape of the pustules 
the two valves described are otherwise identical. 

In conclusion.—The type is being presented to the Tas- 
manian Museum and is figured from a drawing. The co-type 
is figured from a photograph taken by the writer and remains 
in my own collection. In a paper on Polyplacophora of Tas- 
mania, by W. L. May and Dr. Torr (Proc. Roy. Soc. Tas., 
1912, p. 35), a note is made that the valves described above 
were wrongly identified by Hedley and May (Rec. Austr. 
Mus., vol. vii., No. W, 1908) as Acanthochiton crocodilus, 
Torr and Ashby. | 


ACANTHOCHITON SHIRLEYI, Nl. sp. 
‘ Specimen No. 1. Type. 

General appearance.—Broad, girdle wide, densely and 
coarsely spiculose, shell flat and low, the spicules of the girdle 
standing up above the shell. 

Colour.—Some valves are darkhorn-colour, others are 
creamy-white; the dorsal area is creamy-white in some valves. 

Anterior valve.—I cannot notice any rays or undula- 
tions. The sculpture on this valve is largely eroded, but it 1s 
evidently well covered with circular pustules. Insertion plate 
long, slits 5, notch very short but continued on upper side in 


14 


a broad groove to the tegmentum, colour white tinged with 
blue. Measures 34 x34 mm. 

Posterior valve.—Tegmentum very small, about two- 
fifths of total width of valve, anterior portion semi-circular, 
mucro posterior, dorsal area rather narrow, wedge-shaped, 
rugose; the wrinkles following the growth-lines are continued 
across the anterior portion of the dorsal area. The shell 
behind the mucro is very flat, and shows little sculpture; such 
as there is, is composed of shallow broken wrinkles following 
the growth-lines. The dorsal area and the portion behind 
the mucro creamy-white, side areas dark horn and sculptured 
with rows of irregular, flat, more or less circular pustules, 
following the growth-lines and, in places, especially towards 
the outer margin, coalescing. Insertion plate long and very 
broad, slit 11, sutural laminae small and produced forward 
with a deep inward bent between them and the wings of the 
insertion plates. This feature is very marked. A. bednalli 
has a somewhat similar inward fold at the slits, but this 
species has this fold on the opposite side of the wing of the 
insertion plate. Sinus broad, the semi-circular margin of the 
tegmentum only occupies half the width. Colour of articula- 
mentum, pale bluish; the slits are continued half-way to the 
tegmentum in a deep, almost semi-circular groove. This valve 
measures, longitudinally 3 mm., laterally 44 mm. 

Median valve.—The following is a description of valve 
4:—The dorsal area is smooth except for growth-wrinkles 
which are continued from the side areas; this area consists of 
a shallow wedge-shaped depression (this feature is common 
to all valves except valve 2, in which this area is arched; a 
similar variation occurs in a second specimen described here- 
under), some of the growth-wrinkles are in this area broken, 
suggesting an ill-defined string of granules. The lateral and 
pleural areas are not separated but are studded with rows of 
flat circular pustules, following the growth-lines. The slope 
of the sides below the elevated fold that margins the dorsal 
area is slightly curved. Inside, as well as the insertion plates 
and sutural laminae, pale blue, slits 11, sutural laminae 
produced forward, sinus narrow and rounded. This valve 
measures 44 x 44 mm. 

Girdle-—Densely covered with long, coarse, cream- 
coloured spicules; the fringe spicules are a little more slender ; 
sutural tufts long, porcelain-white, and pointed. 

Measurement of dried shell, 15 x6 mm. 

Habitat.—North-west Reef, Barrier Reef, Queensland. 

Remarks.—I am indebted to Dr. John Shirley for the 
opportunity of examining and describing this A canthochiton, 


15 


and I have much pleasure in naming it after him. The speci- 
mens submitted to me were badly encrusted and, in parts, 
eroded; the true characters only became visible on disarticu- 
lation and cleaning. The shape of the articulamentum both 
in median and tail valves, the fact that the dorsal area is 
usually concave, and the girdle densely spiculose, easily 
separates this Acanthochiton from any other Australian 
species. The type belongs to the Queensland Museum. 


Specimen No: 2. Co-type. 


The second specimen, which I am calling the co-type, 
differs from the type in that the centre of the dorsal area, in 
the median valves, is slightly convex, becoming flat or slightly 
concave before the pustulose sculpture is reached. Also the 
pustulose character’ of the sculpture is more limited in area, 
anteriorly the pustules, which are very flat, become sub- 
_ obsolete and confluent following the course of the growth-lines. 
The co-type remains in my collection. 


ACANTHOCHITON RETROJECTUS, Pilsbry, 
var. pustulosus, n. var. 

(A retrojectus, Pils., Naut., vii., p. 107, Jan., 1894; Proc. 
Acad. Nat. Sci. of Phil., 1894.) 

Introduction.—In November, 1918, I collected a very 
long series of A. retrojectus, Pilsbry, in the Quarantine Sta- 
tion, Sydney Harbour. And early in 1919 the following notes 
on this somewhat difficult species were written. At first one 
concluded that there were at least two or three different species 
represented in the series collected from the one spot. Many 
had similar sculpture in the dorsal area to that of the other 
areas, but in some this area was smooth, and, again, while 
many had the regular, evenly-rounded, pustulose sculpture 
described by Dr. Pilsbry; in others a large part of the shell 
was ornamented with large tear-drop pustules. 

On fuller investigation it was found that there was a 
complete series of intermediate forms. All are similar in the 
character and structure of the girdle, the shape and lamina- 
tion and slitting of valves, the tail valve and position of 
mucro; throughout these features seem consistent, and the 
extreme divergence of sculpture does not, in my opinion, 
warrant the separating of them into distinct species. Never- 
theless, at the suggestion of my friend, Mr. W. L. May, I 
propose a distinct varietal name for the form with coarse 
pustules, calling it var. pustulosus. 

Girdle is densely covered with minute scales or short 
blunt-ended spicules, these are mottled, the white ones often 


16 


placed in rings, x65; this ring resolves itself into a string 
of blunt-ended scales, set in a circle. Owing to the minute 
nature of these scales the general appearance of the girdle, 
except under a high power, is spongy. The sutural tufts are 
white and well defined. 

Dorsal area.—Broadly wedge-shaped. In typical shells it 
is ornamented with longitudinal rows of strongly-raised cir- 
cular pustules, set bilaterally in divergent lines, thus forming 
a V with the apex in the centre of the area. But in different 
specimens these pustules vary from circular, well-raised 
pustules to those that are mitre-shaped, or even to long flat 
dashes. Again, in some specimens this area is absolutely 
smooth except in the margins, but this variation appears 
somewhat rare. 

Pleural and lateral areas.—These, in typical specimens, 
are not distinctly differentiated and are ornamented with 
longitudinal rows of strongly-raised circular pustules, gradu- 
ally increasing in size towards the girdle. The varient, which 
I suggest should be known as variety pustulosus, Ashby, has 
the first row or so of pustules from the dorsal area, more or 
less round, but fully half the valve is decorated with a few 
large tear-drop-shaped pustules, some of them being fully 
three or four times as long as wide. In some, the regularity 
of the longitudinal rows is preserved; in others, this system 
of sculpture is lost, these tear-drop pustules being irregularly 
placed, widely separated, and very raised. 

Colour.—While most specimens are mottled pale green 
and black, there are some that are pale green throughout, 
and others that are uniformly rufous; in some the dorsal area 
only is reddish-brown, in others it is pink. When disarticu- 
lated and cleaned the shells are transparent and, usually, both 
tegmentum and articulamentum are green. 

Habitat.—While very numerous at Port Jackson, New 
South Wales, it appears less common in Victoria, and I have 
not taken it in South Australia, where its place is taken by 
the allied form, A. kombert, Torr. This latter I also met with 
in Western Australia. 

Probably the Sydney shell has extended down the east 
coast and then turned towards the west, along the Victorian 
coast; the allied form, A. kimberz, has come in from the west 
and somewhat overlaps A. retrojectus in Victoria. 

I am indebted to Messrs. Gatliff and Gabriel for the 
opportunity of examining a number of specimens from their 
respective collections. 

They were from Western Port, Port Philip Head, Point 
Nepean, Torquay, and San Remo, all in Victoria. They show 


17 


a good deal of variation, mostly of the large pustulose variety. 
Some of the shells were larger than any I secured at Port 
Jackson, and amongst them were certainly some representa- 
tives of A. kimbert. Some forms of A. retrojectus are very 
difficult to separate from that species, unless the specimens are 
very perfect. 

In conclusion.—Dr. Pilsbry, J.c., founded the sub-genus 
Meturoplax for the reception of this species, chiefly on the 
character of the dorsal area, ‘‘dorsal area indistinctly 
differentiated’’; while, in typical specimens, this may be true, 
this feature is not constant, and makes one hesitate to adopt 
his sub-generic name at this stage. ; 


ACANTHOCHITON CORNUTUS, Torr and Ashby, and 
A. ExILIS, Torr and Ashby. 


(Trans. Roy. Soc. S. Austr., vol. xxii., pp. 217-219, Oct., 1898.) 

Until last year A. cornutus was only known from the 
unique type taken by the writer at Marino, in South Aus- 
tralia; but on January 24, 1920, I took a second at Cape 
Jervis. Messrs. Gatliff and Gabriel each lent me a very fine 
specimen that they had identified as A. eaxilis. I noted that 
‘they were con-specific with my type of A. cornutus, and, 
later, compared them with the type of A. ezalis which is in 
Dr. Torr’s collection, and found that A. ezilis is simply the 
juvenile form of cornutus. I find a note in my note-book, 
made a couple of years back, that these two forms were very 
close to one another. ’ 

The largest specimen of the series dredged by Dr. (now 
Sir) Joseph Verco was selected as the type of exilis, and was 
only 3 mm. long; all the specimens were much curled and 
somewhat bleached, whereas the type of cornutus was over 
10 mm. long and well preserved. 

There are slight differences between the two, but not 
more than can be attributed to immaturity; the minute 
curled extlis certainly looked very different from the fine 
specimen of cornutus, but they are undoubtedly the same 
species. As cornutus was described on an earlier page than 
extlis, it has that priority, and A. ezilis is a synonym thereof. 

In the addendum to our paper, lJ.c., reference is made 
to apparent ‘‘eyes’’ on the dorsal area of A. cornutus. It is 
interesting to note that small black specks are visible in the 
shells of the three recently discovered specimens, before 
referred to; but I have not yet been able to determine whether 
they are true eyes, or some other sense organ. The deter- 
mination of their true character must be left to future 
investigation. 


. 18 


ACANTHOCHITON cOxI, Pilsbry. 


(Naut., vii., p. 119, Feb., 1894; Proc. Acad. Nat. Sci. Phil., 
1894, p. 80, pl. ili, , figs. "21-26, pl. iv., fig. 834; A. lachrymosa, May 
aud pe P. and Proc. Roy. Soc. Tas. , 1912, pp. 36 and 37, pl. i., 

gs. 1-4 

The identification of this Acanthochiton has always been 
a difficulty with me. In July, 1919, I wrote the Australian 
Museum for the loan of a specimen, and at their suggestion 
Mr. Bassett Hull very kindly sent me a shell which, on exam- 
ination, I found could not possibly be A. coai, but, strangely 
enough, it was a worn specimen of a species described by 
Dr. Torr and the writer in October, 1898, under the name 
of A. crocodilus, and which, up till the identification of Mr. 
Hull’s specimen from New South Wales, was only known to 
occur in South Australia, and limited to the pair originally 
described, which were side by side on the same rock, at low 
water, at Marino. 

Later on I received specimens from Dr. Torr’s collection 
and the Queensland Museum, labelled A. coz, but in both 
cases they were misidentifications and referable to well-known 
species. Through some oversight, although I again applied 
to the Australian Museum for the loan of their co-type, it 
was never sent for my inspection. | 

In correspondence with Professor Dr. J. Thiele, of Berlin, 
I mentioned my desire to see coat, and he was good enough 
to send me a specimen from Balmoral, which I conclude is the 
place of that name in North Borneo. The specimen was 
marked “‘identification uncertain.’’ If it had come from Tas- 
mania one would have no hesitation in identifying it as a 
slight variant of A. lachrymosa, May and Torr. 


It differs from the Tasmanian shells, slightly, in the 
arrangement of the pustules, and the spicules on the girdle 
are slightly coarser. On writing Mr. W. L. May he advised 
me that he had some years ago seen the co-type of cozz in the 
Australian Museum and had made a note that it was very 
close to Jachrymosa. In October last, Mr. May brought over 
a very fine series of lachrymosa from the type locality. We 
found that the sculpture varied from long, slender, flat, finger- 
like processes to short oval discs, or elongated tear-drop 
pustules. Most in the juvenile stage have quite small pus- 
tules, but even in this they are not consistent. They also 
vary very much in the spacing of the pustules; mostly they are 
crowded, as is so well shown in the figures accompanying May 
and Torr’s description, J.c. 

A comparison of these figures with the figures in Dr. 
Pilsbry’s paper on ‘‘Port Jackson Chitons,’’ /.c., will explain 
the difficulty we have all laboured under in identifying 4. 


19 
coxt, Pilsbry. The description in Pilsbry’s paper would do — 


for lachrymosa. 


In conclusion.—While all previous records of A. lachry- 
mosa have been confined to about 150 yards of beach, on 
Frederick Henry Bay, southern Tasmania, the writer was 
able some time ago to extend its range to Sulphur Creek, 
north-western Tasmania; and, more recently, Mr. May has 
found it on Bruny Island, and now Mr. May and myself are 
satisfied that it is con-specific with A. cozi, Pilsbry. . We 
have the record of two specimens taken by the late Dr. Cox, 
at Port Hacking, New South Wales, and, finally, Dr. Thiele’s 
specimen extends its range far into the tropics. 

Thus a species that has hitherto been considered one 
of the most restricted in its range is found to have a most 
extended range north and south, probably greater than any 
other of our known specimens. It certainly appears very local 
in its occurrence, the reasons for which must await further. 
elucidation. I have sent two specimens to Dr. Pilsbry, asking 


-him to kindly compare with his type of A. coxi,.and advise 


whether he can find any justification for retaining the Tas- 
manian shell as a sub-species of A. cozt, Pilsbry. 
Notr.—Since the foregoing was written I have received 
Dr. Pilsbry’s reply, which is as follows:—“‘‘I have carefully 
compared the specimens of A. lachrymosa, May and Torr, 
with the type of A. coz. I am satisfied that there is no 
specific difference. A sub-specific difference may be indicated 
by (1) the difference in colour, my form being pink within, 
yours greenish; (2) the wider central areas of valves 3-8 in 


my specimen. This is exaggerated in the figures, which were 


done on stone by a commercial lithographer from my pencil 
drawings.”’ 

This fully confirms our opinion, and in face of the 
variability of this species we are hardly justified in making 
a sub-species of the southern form. A. lachrymosa, May and 
Torr, is therefore a synonym of A. cozz, Pilsbry. 


The pitting of CaLLocHITON PLATESSA, Gould, var. fossa, nov. 
(Proc. Bost. Soc. N.H., ii., 1846, p. 143.) 


’ Some years ago I noted that one of the shells belonging 
to this species, that I had collected in Gulf St. Vincent, 
showed six deep pits, immediately in front of the lateral area 
of the seventh valve. On January 24, 1920, I collected a 
second specimen in which the same valve shows a similar 
number of pits. A few months back, when going through 
the collection of the Polyplacophora in the South Australian 
Museum, with a view to determining the species, I noticed a 


20 


similar specimen with pitting confined to the same valve; all 
these were taken in this State. In going through a score of 
specimens in my own collection from New South Wales, © 
Tasmania, South Australia, Western Australia, and New 
Zealand, with the exception of one specimen 25 mm. long 
from Sydney, which is pitted in the seventh valve, all are 
typical unpitted shells. Mr. W. L. May has sent me for 
examination three large and handsome specimens from Watson 
Bay, New South Wales, all of which show similar pitting on _ 
the seventh and eighth valves, and the largest one, which is 
over 40 mm. long, has incipient pitting in the sixth valve 
as well. In this specimen I counted 12 pits in the seventh 
valve. These pits commence high on the ridge in the juvenile 
shell. The pits are deep and only a little longer than broad, 
in fact very similar to the pits near the ridge of Rhyssoplax 
oruktos, Maughan, but there the likeness ends—they are not 
as regular in shape nor developed to the same length as in 
that species. Again, the character of the pitting is quite 
different from C. rufus, Ashby. I have compared it with » 
the type and with the juvenile form from the Bracebridge 
Wilson collection ; the grooving of rufus can hardly be termed 
pitting, but is really longitudinal grooving, and is present to 
an equal extent in all the valves except the first. 

In concluston.—The existence of these pits and their 
occurrence consistently on the seventh valve, and in the case 
of the Watson Bay specimens.on the eighth valve as well, 
suggests a definite tendency to vary in this direction. At first 
I thought of suggesting that deep-water specimens may have 
a greater tendency to develop this form of sculpture, and that 
C. rufus (which is only known from dredge specimens) may 
have been derived from such a pitted race of C. platessa. 
On more careful examination, however, I do not feel justified 
in advancing such a.hypothesis. It will be well worth while 
for collectors to keep their eyes open for this variant, which 
may well be known as Callochiton platessa, var. fossa, Ashby. 


SYPHAROCHITON PELLIS-SERPENTIS, Quoy and Gaimard, 1835. 


(Chiton pellis-serpentis, Quoy and G. Voy, Astrol., ii., 381, 
pl. 74, f. 17-22; Man. Conch. (1), xiv., 173,-pl. 37, f. 14-17; Bee 
Mal. Soc., ii., 195, c. Squamosus, L. Wissel, Zool. Jahrb., xx., 
619, not of Linne (Anatomy); Tate and May, Proc. Linn. Soc. 
N.S. Wales, 1901, pt. 3, pp. 412-415; May and Torr, P. and Proc. 
Roy. Soc. Tas., 1912, pp. 38 and 39. Type, Mus. Hist., Paris.) 


SYPHAROCHITON SINCLAIRI, Gray, 1843. 
(Dief., N. Z’d, ii., 263; Man. Conch. (1), xiv., 174,«pl. 36, 
f. 1-3; Proc. Mal. Soc., ii., 196; Wissei, Zool. Jahrb., xx., 627, 
pl. 23, f. 38-44, pl. 24, f. 45-48 (Anatomy). Type, Brit. Mus.) 


21 


SYPHAROCHITON MAUGEANUS, Iredale and May. 
(Proc. Mal. Soc., vol. xii., pts. 11. and iii., p. 114-115, Nov:, 1916.) 

In October, 1921, Mr. W. L. May, of Tasmania, and the 
_ writer jointly examined a fairly large series we had collected 
in different parts of Tasmania with shells in my collection 
from New South Wales and New Zealand. My New Zealand 
specimens were from various localities, but the S.. senclairi, 
Gray, were from Doubtless Bay, collected by Mr. Albert E. 
Brookes, and from Te Onepote, collected by the late Mr. 
Suter. 

We cannot agree with Iredale and May in separating 
the Tasmanian shells from the New Zealand ones, or from 
those from New South Wales. Pilsbry, in his paper on ‘‘Port 
Jackson Chitons’’ (1894), also states that he ‘‘was unable to 
detect any difference between New South Wales and New 
Zealand shells.’’ Therefore S$. maugeanus, Ire. and May, 
becomes a synonym of S. pellis-serpentis, Quoy and Gaimard. 

Further, we find that the smooth shells living in company 
with the more sculptured ones, in Frederick Henry Bay, Tas- 
mania, are con-specific with S. sinclair, Gray, 1843, the New 
Zealand shells being similar to the Tasmanian ones. In both: 
S. pellis-serpentis varies from the somewhat flat highly- 
sculptured shells so common in Port Jackson, New South 
Wales, to those that are almost smooth in all areas. We 
therefore consider that S. senclairi is a smooth variant of S. 
pellis-serpentis, and is certainly common to New Zealand and 
Tasmania, and, on the authority of the late Dr. Cox, we 
must conclude, of New South Wales as well, although neither 
Mr. May nor the writer has seen the smooth variety from New 
South Wales. 

Mr. May sends me the following note in reference to 
the foregoing:—‘“‘I agree with all you have written. The 
shell varies very much in height, some being very flat, others 
high and round backed, with all grades between; they also 
vary greatly in size in different localities. My largest, from 
- Wedge Bay, is 56 mm. long; they may be almost white, black, 
or of varying patterns of black and white, etc.”’ 

In conclusion.—We find that Sypharochiton  pellis- 
serpentis, Quoy and Gaimard, is an extremely variable shell 
in Tasmania, New Zealand, and New South Wales, varying 
from a highly-sculptured form to an almost smooth one, which 
must be known as variety sinclairi, Gray, the intermediates 
still living ; and Iredale and May’s S. maugeanus is a synonym 
of S. pellis-serpentis, Quoy and Gaimard. 


22 


LorIcELLA ANGASI, H. Adams and Angas. 
(Proc. Zool. Soc., 1864, p. 193.) 


In my paper on the above genus (Trans. Roy. Soc. 8S. 
Austr., vol. xliii., 1919, pp. 59-65) reference is made to the 
‘‘finger-like processes’’ being noticeable in the anterior portion 
of the girdle, and the ‘‘spear-head spicules’ being set opposite 
them and apparently having some relation thereto. in 
January, 1920, I collected a very well-preserved specimen at 
Marino, South Australia, which was free from the usual 
foreign growth. In this specimen, which is 60 mm. in 
length, the finger-like. processes of the girdle extend right 
round, and the remarkable ‘‘spear-head spicules’’ are placed 
opposite these, right round the girdle. In a letter, dated 
October 17, 1921, Mr. S. Stillman Berry, of Redlands, Cali- 
fornia, writes me in answer to a letter of mine referring to 
some remarks that had been made in reference to these strange 
spicules on Loricella:—‘‘I have worked on those Loricella and 
Komonella spicules just enough to know that I want to go 
into the matter of their structure a great deal more 
meticulously, which will mean a lot of work in the preparation 
of slides and so on. I cannot understand how anyone can 
interpret chiton setae as algae.’’ 


DESCRIPTION OF PLATE III. ; 


Fig. la. Acanthochiton mayi, Ashby, portion of median valve. 


Type. 
rm, “i 4 », median valve. Co-type. 
x abt. 13 times. 
Sp i shirleyi, Ashby, portion of posterior valve. 
Type. 
5° 2203 59 iN » portion of median valve. 
Type. 
A a2e. 3 3 » median valve. Co-type. 
x abt. 11 times. 
sien aoe is gabrieli, Ashby, portion of median valves 


showing longitudinal striae in dorsal 
area. Type. 
» 4. Callochiton platessa, var. fossa, Ashby, portion of 7th 
valve showing pits. 


23 


A NEW ISOPOD FROM CENTRAL AUSTRALIA BELONGING 
TO THE PHREATOICIDAE. 


By Cuaries Cuitton, M.A., D.Sc., C.M.Z.S., 
Professor of Biology, Canterbury College, New Zealand. 


(Communicated by Professor F. Wood Jones). 
[Read April 13, 1922.]| 


In this paper I describe a new and most interesting 
fresh-water Isopod kindly sent to me by Professor F. Wood 
Jones, of Adelaide University. It was collected in June, 
1920, in artesian water from the Hergott (Marree) bore, in 
Central Australia, a little south of Lake Eyre. 


The animal proves to belong to the Phreatoicidae and 
comes sufficiently near the typical genus Phreatoscus to be 
placed in it. The Phreatoicidae is a family of fresh-water 
Isopods of which the first member was described in 1883, from 
the underground waters of the Canterbury Plains in New 
Zealand. Later on other species of the genus, and of closely 
allied genera, were described from the surface and under- 
ground waters of Australia, and, still more recently, Barnard 
(1914, p. 231) recorded a species of Phreatoicus from the 
‘mountain streams of Cape Colony, South Africa. The family 
is quite distinct from all the other families of the Isopoda, 
and forms, by itself, the sub-order Phreatoicidae, marked by 
some primitive characters and by a striking but superficial 
resemblance to the Amphipoda. The characters and distri- 
bution showed that the family must be an ancient one, and 
in 1918 this was proved by the discovery of a fossil species 
from the Triassic beds of New South Wales. The fossil form 
is not very different from some of the existing species, and, 
apparently, members of the family have been living in fresh 
waters on some part of the Australian continent from Triassic 
times up to the present. The discovery of another quite dis- 
tinct species in Central Australia is most interesting and 
important as confirming the conclusions already arrived at. 
Further details of the history of the family will be found in 
my paper describing the fossil species (Proc. Roy. Soc. N.S. 
Wales, vol. 51, p. 383). 

The mode of occurrence of the new species is worthy of 
note. In his first letter, Professor Wood Jones said :— 
‘‘Hergott is a pure artesian bore; the water is hot, and the 
creatures were in thousands swimming in the hot water near 
the bore head.’”’ The specimens sent were found to possess 
well-developed eyes and to be of a dark-slaty colour, so that 


24 


they evidently had not come up the bore from underground 
waters. On my pointing this out and asking for further par- 
ticulars, Professor Wood Jones wrote: —‘‘Now I have asked 
everyone who knows, and I am assured that all the water is 
bore water pure and simple.. At Hergott there are natural 
springs—that is why the place sprang into existence. I have 
never seen the springs; they are some three miles away from 
the place where the bore was sunk. . . . The bore is 
just on the desert—the water flows on the desert where pre- 
viously no water was (there is no old watercourse into which 
the bore water has found its way, as at Clayton and Dul- 
canina).’’ It is no wonder, therefore, that it is the popular 
belief that the animals came up the bore, for this is, as 
Professor Wood Jones says, ‘‘the local story of all bore-water 
fauna.’’ He adds that it is curious that though every party 
that has gone into the centre of Australia has based on 
Hergott, no one has noticed or collected the Isopod, although 
the hot water of the bore is full of them. When he was there 
they were in countless numbers, all swimming against the 
hot current. He did not take the temperature of the water, 
but says, ‘“‘It is very hot; steam arising from it.”’ 

Like other Isopods, the Phreatoicus carries its eggs in a 
brood pouch underneath the body till the young are 
hatched out and, probably, for some time longer, the young 
then being similar in form to the adults. It is, therefore, a 
little difficult to see how they have got from the spring, or 
other natural water from which they must have come, to the 
bore water in which they exist in such numbers. It is, of 
course, possible that when the natural water dries up they 
become encased in the dried-up mud, retaining the power 
of vitality and resuming activity as soon as the water reappears, 
but that does not explain how they have got from the natural 
springs, situated near Marree, to the bore water, three miles 
distant. It is, however, clear that they must be widely dis- 
tributed and abundant in springs and natural waters in the 
district, for Professor Wood Jones, in a letter dated October 
5, 1921, states that in a recent trip he collected specimens 
from the mound springs, near Coward, just to the westward 
of Lake Eyre south. There are, he says, many of these 
springs, and they vary greatly in salinity and temperature, 
but the animal was found in all the springs, from Bullakaninna 
to Coward, an area of some 30 miles. 

In this connection it is worthy of note that another mem- 
ber of the family, Phreatoicopsis terricola, Spencer and Hall, 
was found in burrows on the banks of the Upper Gellibrand 
River (Spencer and Hall, 1896, p. 13). This species has since 
been recorded from the Otway forest; from Mount William, 


25 


near Ararat; and from the Grampians (Raff, 1912, p. 70). 
Another species, Hypsimetopus intrusor, Sayce, occurs in the 


burrows of the land crayfish, Engaeus cuniceularius, in Tas- 


mania (Sayce, 1902, p. 218). The remaining species of the 


e 


family appear to be genuinely aquatic, being found in surface 
or underground fresh-water streams. 

Although the species under consideration is being placed 
for the present under the genus Phreatorcus, it differs from 
the other members of the genus in at least two characters. 
The more evident of these, though not the more important, 
is the greater expansion of the basal joints of the last three 
pairs of peraeopoda, as shown in figs. 1 and 10. In the other 
species of the genus these joints are comparatively narrow, 
as in most Isopods™; but in the present species the ex- 
pansion is fully as great as that in most Amphipoda, and still 
further increases the resemblance to an Amphipod, caused from 
the laterally compressed form of the body. It may be men- 
tioned, however, that the next joint, the ischium, is com- 
paratively long—longer than the succeeding joint, the merus— 
while, as I have elsewhere pointed out (1894, p. 205), in 
Amphipoda, with broadened basal joints, the ischium is 
usually quite short. 

The other point of difference, though less evident, is of 
more real importance, viz., the apparent absence of the coxal 
joints of all the peraeopoda. In other species this coxal joint, 
though small, is quite well marked and can be readily recog- 
nized as the first joint of the limb, for it is not flattened 
into a side plate or “‘epimeron,’’ as it is in most Amphipoda. 
In P. latipes the pleura of the first four segments are pro- 
duced downwards and outwards so as to hide the base of the 
leg, and even when the attachment of the limb to the inner 
side of the pleuron is examined, nothing is seen that can be 
definitely recognized as the coxal joint. Consequently it must 
either have become fused with the pleuron, but if so without 
any suture or mark indicating its presence, or it is quite 
absent. Calman (1909, p. 202) has some interesting remarks 
on the development of the coxal joint of the peraeopoda in 
various Isopods, and gives examples in which it appears to 
replace the pleural expansion of the segment, though, in 
that case, it is marked off from the segment on the dorsal 
surface by a distinct suture, except in the first segment, where 
there is no suture, and in some of the Oniscoidea in which 
the suture on the other segments also may disappear. 


() The basal joints are slightly broadened in ‘Phreatoicus 
australis. 


B 


26 


The species of Phreatovcus now under consideration may 
be described as follows : — 


PHREATOICUS LATIPES, 0. sp. 
Figs. 1-14. 

Specific diagnosis.—Body stout. Peraeon (fig. 2) broad, 
not laterally compressed, moderately convex with the pleural | 
portion of the first four segments projecting outwards and 
slightly downwards so as to conceal the basal joints of the legs. 
Pleon short, about half the combined length of the cephalon 
and peraeon, moderately compressed laterally, pleural portions 
of the segments produced downwards, their lower margins 
being rounded and fringed with a few setae. First segment 
of peraeon short and immovably joined with the head but with 
the suture well marked, pleural portion of segment free and 
produced anteriorly about half-way along the lower margin of 
the head, those of the second and third segments less pro- 
duced anteriorly. Hye well developed, irregularly rounded or 
subtriangular, black. Surface of the body covered with small 


scattered setae, nearly smooth but with slight wrinkles or 


irregularities on most of the segments. Sixth segment of 
peraeon United with the terminal segment, or telson, but 
distinctly marked off from it by a well-defined suture running 
obliquely backwards from the upper pleural portion of the 
fifth segment to the base of the uropod (fig. 4). Terminal 
segment strongly arched above, sides widely separate below, 
the mid-dorsal end portion showing as a slight process in side 
view and when seen from above having a median indentation 
between two rounded lobes, each of which bears three or four 
setules. (Fig. 3.) 

First antenna more than half the length of the second, 
joints of the flagellum not broadened. Second antenna nearly 
as long as the head and first two segments of the peraeon. 


The mouth parts do not differ greatly from those of 
Phreatoicus australis. In the mandibles the palp is rather 
short, the third joint being quite short and bent at right 
angles to the second. There are two strongly chitinized cut- 
ting edges in the left mandible; in the right the inner one is 
small and colourless, as in P. capensis. The first. maxilla has 
about six plumose setae at the apex of the inner lobe. In the 
second maxilla the two outer lobes are very slender, bearing 
long pectinate setae; the inner lobe is broader and rounded, 
densely setose, and fringed along its inner margin with a very 
regular and distinct row of long setae. In the maxilliped, 
the epipod is nearly circular, thin; the second joint bears a 
very distinct row of plumose setae projecting inwards towards 
the mouth cavity ; the palp is of the usual structure. 


Fic.2 FIGs 


All the figures refer to Phreatoicus latipes and are taken 
from a male specimen. 
Fig. 1. Side view of the whole animal. 
P- >», 2 Dorsal view of body. 
,, 3 Dorsal view of end portion of terminal segment. 
; ,, 4. Side view of pleon, straightened out to show the 
anterior segments more clearly. 


28 


First pair of legs strongly subchelate; second and third 
similar to one another, feebly subchelate; fourth pair more. 
slender and not specially modified in the male; fifth, sixth, 
and seventh pairs increasing progressively in length, their _ 
basal joints: flat and greatly produced posteriorly into a 
rounded lobe similar to that in many Amphipoda, the lobe 
marked off from the joint proper by a distinct ridge, posterior 
margin of the lobe entire (fig. 10). 

Uropods short, not projecting much beyond the end of 
the terminal segment, outer branch slightly shorter than the 
inner. 


Colour.—Dark slaty-grey. In some young specimens the ~ 
surface of the body is lighter in colour with dark pigmented 
spots much more widely separated from one another than in 
the adult. 

Length of body (in curved position), about 15 mm. 
Greatest breadth of peraeon, about 6°5 mm. 


Locality.—In hot water from Marree (Hergott) bore, 
and in springs and streams near Coward, Central Australia. 
Collected by Professor F. Wood Jones, Adelaide University. 


Remarks.—Although in the flattened character of the 
peraeon and the greatly broadened basal joints of the last 
three pairs of legs this species differs markedly from other 
species of Phreatoicus, there seems to be a fairly close 
resemblance in the various appendages, so that it will not 
be necessary to give a very detailed account of these. 


The first antenna (fig. 5) is slender, the first and third 
joints of the peduncle similar and considerably longer than 
the second; the flagellum is about the same length as the 
peduncle and contains about ten joints, which bear short 
simple setae and a few olfactory setae. The second antenna 
(fig. 6) is considerably longer and stouter than the first; 
the first two joints of the peduncle are short, the third 
about twice as long as the second and subequal with the 
fourth, the fifth longer and more slender; the flagellum is 
subequal in length with the peduncle and contains about 
nineteen joints, the basal ones being somewhat stout and bear- — 
ing tufts of numerous short simple setae. 


In the male the legs of the first pair (fig. 7) are strongly 
subchelate, the propod being subtriangular and greatly 
broadened at the base, the finger not reaching beyond the 
straight palm. In general appearance this appendage is 
similar to that of P. australis. The second and third pairs 
of legs (fig. 8) are similar, longer, and more slender than the 


All the figures refer to Phreatoicus latipes and are taken 
from a male specimen. 


ig. 5. 


6. 
& 
8. 
o: 


First antenna. 
Second antenna. 
First peraeopod. 
Third peraeopod. 
Fourth peraeopod. 


30 


first; the propod is not broadened, but the finger is very 
long, slightly curved, and when flexed reaches back as far as 
the basal portion of the carpus, forming apparently an efficient 
grasping organ. The fourth leg (fig. 9) is slightly longer 
than the third with the joints more slender, and it is not sub- 
chelate but simple, the finger not longer than the propod. 
This appendage is the same in both male and female, although 
in some other species of Phreatoicus the legs of the fourth 
pair are modified in the male to form a special grasping organ. 
The fifth, sixth, and seventh pairs are quite similar, increasing 
progressively in length posteriorly. The basal joint in each 
is very greatly expanded behind into a rounded lobe pro- 
jecting backwards and downwards, reaching two-thirds of 
the way to the distal end of the ischium. This expansion 
is marked off from the joint proper by a distinct ridge running 
parallel to the anterior margin; the posterior margin of the 
lobe is entire and bears no setae; the ischium is distinctly 
longer than the merus and, like it, broadened somewhat 
distally ; the carpus and propod are cylindrical; the finger is 
straight, acute; these joints show setae of varying sizes, as 
indicated in the figure (fig. 10). 


The male appendages (fig. 11) on the seventh peraeon 
segment are slender, tapering, curved inwards towards one 
another, slightly swollen at the base, and apparently grooved 
on the posterior surface. 


The pleopods show a close general resemblance to those 
of P. australis. The first pleopod has the basal joint or 
protopod short, the endopod and exopod subequal, each form- 
ing an irregular oval lobe, the margin of the endopod being 
smooth and without setae, as in all the pleopods, the outer 
margin and apex of the exopod being fringed with fine setae. 
The second pleopod in the male (fig. 13) has the basal joint 
broader and bearing a few long setae at its inner margin; 
the endopod is similar to that of the first pleopod, but bears 
on the inner side the penial appendage, which is four-fifths 
as long as the exopod, broadened near the base and apparently © 
grooved on its upper or anterior surface; the exopod is larger 
than the endopod and consists of two joints, the basal one 
about as long as the endopod and produced at its outer 
proximal angle into a broad rounded lobe; the terminal 
joint is small, oval, and has its margins fringed with long 
setae, a few long setae being also present on the distal portion 
of the outer margin of the basal joint. The third (fig. 14), 
fourth, and fifth pleopods are similar to the second, except 
for the absence of the penial appendage, and they all bear 
attached to the outer margin of the basal joint a large 


All the figures refer to Phreatoicus latipes and are taken 


Fig. 


23 
2) 
be) 
3 


from a male specimen. 


Seventh peraeopod (less highly magnified than 
figs. 7, 8, and 9). 

Male appendage. 

First pleopod of male. 

Second pleopod of male. 


_ Third pleopod of male. 


32 


well-developed oval “‘epipod,’’ the margins of which are 
fringed with long setae. 

The uropods are similar to those of other species of © 
Phreatoicus, having the basal joint subequal in length with 


the branches, its upper margin fringed with stout setae, the 


upper margin of each branch being similarly fringed. 

A ffinitces.— Until it is possible to make a revision of the 
Phreatoicidea this species may be left’ under the genus 
Phreatoicus. It shows a good general resemblance to P. 
australis, but differs markedly from that species, and indeed 
from all the members of the tribe, in the absence of the coxal 
joints of the peraeopoda. It resembles P. australis in having 
the first peraeon segment short and more or less fused with 
the head, in this character agreeing also with Phreatoicopsis 
terricola, Spencer and Hall. It agrees with the latter species 
and differs from Phreatorcus australis in the fact that the 
fourth peraeopod is not specially modified in the male. The 
sixth segment of the pleon, although fused with the terminal 


segment, or telson, appears to be more distinctly marked 


off from it by a distinct suture than in the other species; in 
Phreatoicus australis there is a suture present, but this ex- 
tends anteriorly only a short distance from the base of the 
uropod and does not reach the posterior margin of the fifth — 
segment. In most Isopods, except the Anthuridae, the sixth 
segment is completely fused with the telson without any 
apparent suture to indicate the line of juncture. 

I am greatly indebted to my assistant, Miss E. M.. 
Herriott, M.A., for preparing the drawings for this paper, 
and to ‘Professor F. Wood Jones for the opportunity of 
describing this interesting species. 


List oF REFERENCES. 
Barnard, Keppel H. 


1914—1. Contributions to the Crustacean Fauna of South 
Africa. 2. Description of a New Species of 
Phreatoicus (Isopoda) from South Africa. Ann. 
South African Mus., vol. x, pp. 231-240, pls. 23 
and 24. 

Calman, T. W. 


1909—Crustacea, in Ray Lankester’s Treatise on Lor IBY 
part vu., Appendiculata, 3rd Fascicle. 
Chilton, C. 


1894—The Subterranean Crustacea of New Zealand, with 
some general remarks on the Fauna of Caves and 


Wells. Trans. Linn. Soc., Zool., vol. 6, pp. 
163-284, pls. 16-23. 


33 


1918—A fossil Isopod belonging to the fresh-water genus 
Phreatoicus. Proc. Roy. Soc. N.S. Wales, vol. 51, 
; pp. 365-388, with text figures. 
Raff, Janet W. 
1912—Notes on the Isopod, Phreatoicopsis terricola, Spencer 
and Hall. Victorian Naturalist, vol. 29, pp. 70-72. 
Sayce, O. A. 
1902—A new genus of Phreatoicidae. Proc. Roy. Soc. 
Vict., vol. 14, pp. 218-224, pls. 18 and 19. 


Spencer, B., and Hall, T. 8. 


1896—Description of a new genus of Terrestrial Isopoda 
: allied to the genus Phreatoicus. Proc. Roy. Soc. 
Vict., vol. 9, pp. 12-21, pls. 3 and 4. 


34 


THE FLORA AND FAUNA OF NuyT’sS ARCHIPELAGO AND 
THE INVESTIGATOR GROUP. 


NO. 1-THE AMPHIPODA AND ISOPODA. 


By Cuar.es Cuitton, M.A., D.Sc., C.M.Z.S., 
Professor of Biology, Canterbury College, New Zealand. 


(Communicated by Professor F. Wood Jones.) 


[Read April 13, 1922.] 


The crustacea referred to in this short paper were collected 
by Professor F. Wood Jones, of Adelaide University, in an 
expedition made towards the end of 1920 to the Investigator 
Group and the Nuyt’s Archipelago, lying to the west of Eyre 
Peninsula, South Australia. The whole of the species here 
mentioned were, however, obtained at the Nuyt’s Archipelago, 
most of them in Smoky Bay. They are all referred to species 
already known, though in some cases they have not been 
hitherto recorded from South Australia. 


In addition to these species numerous specimens of 
terrestrial Isopods, belonging to Cubaris and allied genera, 
were collected from several localities. These have been sent 
for determination to Dr. W. E. Collinge, York Museum, 
England. Several shore Amphipoda (Orchestia, etc.) were 
obtained at various places, but they are all too immature for 
determination. 


AMPHIPODA. 
LEUCOTHOE SPINICARPA (Abildg.). 


Tebeniiee spinicarpa and L. miersi, Stebbing, 1906, p..165. 


Leucothoe commensalis, LL, diemenensis, and L. gracilis, 
Stebbing, 1910, p. 636. 


Leucothoe spinicarpa, Chilton, 1912, p. 478, and 1921, p. 59; 
Barnard, 1916, p. 148. 

Locality.—Smoky Bay, South Australia, 3°5 to 4 fms. 
Two specimens; length, 10 mm. 

In my report on the Amphipoda collected by the F.I.S. 
‘‘Endeavour,’ I have given reasons for considering all the 
forms mentioned above as belonging to the cosmopolitan 
species, L. smnicarpa (Abildg.). The species seems to be com- 
mon at numerous places on the Australian coasts. Barnard - 
has given some further particulars of specimens from South 
Africa, and has added L. muiersi, Stebbing, to the list of 
synonyms, as I had already done in my MS. notes. 


— 35 


GRUBIA SETOSA (Haswell). 

Amphithoé setosa, Haswell, 1879, p. 270; Chilton, 1885, 
p. 1040. 
Grubia setosa, Stebbing, 1906, p. 644, and 1910, °p. 649. 
Locality.—Mangrove Creek, Smoky Bay. Several 
specimens. 

I refer these specimens to the species named with some 
doubt, for they are all small and immature, and the species 
itself is imperfectly known. The typical species of the genus, 
G. crassicorms, is known from the Mediterranean and the 
Black Sea, and a South African one has been described by 
Barnard under the name G. australis. It will be necessary 
to compare adult specimens of these three species before any- 
thing can be said about their affinities. 


ISOPODA. 


Dero marina (Chilton). 


Deto marina, Chilton, 1915, p. 444, pl. 39, figs. 19-23. 

Deto marina, Chilton, 1917, p. 399, figs. 15-21. 

Localities.—Smoky Bay, 21-xi.-20. Two specimens. 
Laura Bay, 23-xi.-20. Five specimens. Eyre Island, Smoky 
Bay, 21-xi.-20, Several specimens. Unnamed guano island, 
Laura Bay, 22-xi.-20. Two specimens. 

This species was originally described under the name 
Philougria marina from specimens collected at Coogee, New 
South Wales, in 1884. No further specimens were obtained 
from the type locality until towards the end of 1920, when 
several were obtained by Mr. F. A. McNeill, of the Aus- 
tralian Museum, Sydney. It had been collected at Kangaroo 
Island, South Australia, by W. H. Baker, in 1915; and I have 
since had specimens from Tasmania, collected by A. M. Lea, 
of the Adelaide Museum. Apparently it is fairly common 
in the localities examined by Professor F. Wood Jones, and 
was obtained at the four places mentioned above. The speci- 
mens agree closely with the description given of those from 
Kangaroo Island. . 

Mr. F. A. McNeill, who collected the specimens from 
Coogee, states that they were found on the damp under- 
surfaces of stones which formed heaped accumulations of small 
sandstone boulders at highest tide mark,and ended among the 
dark crevices and overhanging shelves of larger rocks, from 
10 to 15 ft. further back. He further states that the animals 
are ‘“‘slow in movement, often lying motionless in the 
irregularities on the surface of the stones; the older examples 
rarely move away until disturbed previous to capture,’’ and 
he contrasts their slow movement with the active movements 
of Ligia australiensis, which was found at the same time and 


36 


place, and was so active that it was very difficult to capture. 
I had noticed the same characteristic habits in the species 
Deto bucculenta, Nicolet, found on the shores of Paterson 
Inlet, Stewart Island, New Zealand (1917, p. 404). 

In my paper on the genus (1915) I have drawn attention 
to the distribution of the different species on islands and 
other land masses in southern seas. 


PaRIDOTEA UNGULATA (Pallas). 


Idotea ungulata, Miers, 1881, p. 52; Chilton, 1890, p. 196. 
lias ungulata,. Stebbing, 1900, p. 538; Collinge, 1918, 
p. 8l. 

Locality.—Mangrove Creek, Smoky Bay. Four speci- 
mens; length of largest, 40 mm. : 

Colour (in spirit).—Olive-green with lighter patches some- 
what irregularly arranged in longitudinal rows. 

These specimens agree, generally, with New Zealand 
specimens referred to this species, though the colour is a little 
different, and the first segment of the pleon seems rather more 
distinct and slightly longer in the median line; in the New 
Zealand specimens this segment is less distinctly marked and 
in the median line is nearly concealed beneath the last seg- 
ment of the peraeon. Collinge (1918, p. 82) has established 
a new variety, atrovirens, for specimens from Victoria, Aus- 
tralia, having the ‘‘whole of the body a very dark olive-green, 
almost black.’”’ From the details given by Miers and by 
Stebbing the colour appears to vary considerably in this 
species. Most of the New Zealand specimens that I have 
been able to examine in the living condition are a light green, 
corresponding with the colour of the green seaweeds on which 
they are usually found. This colour disappears in spirit 
specimens, leaving them a yellowish-brown. Some of my 
specimens, however, still have (in spirit) the whole body more 
or less darkly coloured ; sometimes the whole body, sometimes 
certain portions only, being finely dotted with black. 

The mouth parts have been described by Stebbing, and 
also by Collinge, the two descriptions showing considerable 
differences. I have a slide with the-mouth parts of a small 
New Zealand specimen mounted about the year 1890. In it 
the first maxilla has the inner plate narrower than in Col- 
linge’s figure and with: only three plumose setae at its 
extremity. Collinge found four and Stebbing (apparently 
describing South African specimens) found ten; the outer 
lobe of this maxilla bears about ten stout spines with one or 
two more slender ones agreeing on the whole with Collinge’s 
figure, though the arrangement differs a little in detail. The 
maxilliped agrees pretty closely in general shape with the 
figure given by Collinge, but the parts corresponding to the 


37 


second and third joints of the palp, as described -by him, are 
almost completely fused, the suture between them being very 
indistinct compared with the articulations of the other seg- 
ments ; consequently the palp appears four- “Jointed as described 
by Stebbing. 

Distribution.—The species is very widely distributed in 
southern seas. 

Idotea excavata, Haswell, comes near to this species, and 
I referred it to Paridotea ungulata in 1890, though, at the 
same time, pointing out several slight differences. 


CyMODOCE LONGICAUDATA, Baker. 
Cymodoce longicaudata, Baker, 1908, p. 188, pl. ii., figs. 1-11. 
Locality.— Mangrove Creek, Smoky Bay. Four specimens. 


These specimens agree well with Baker’s description, 
though in the largest (length of body with terminal spine, 
15 mm.) the terminal spine, the branches of the uropoda, and 
the side-plates of the peraeon are longer and more acutely 
produced than in his figure. 

A specimen of this species has recently been sent to me 
by Professor F. Wood Jones, labelled ‘‘Onkaparinga River, 
Mt. Lofty,’’ presumably in fresh water. 


ZUZARA VENOSA (Stebbing). 

Zuzara venosa, Baker, 1910, Trans. Roy. Soc. S. Austr., 
vol. xxxiv., p. 83, pl. XXiii., figs. 13-16, and pl. xxiv., figs. 1-3. 

Several specimens, es on oe shore of onal Bay, 
South Australia. 

Of these, three are fully adult males; the others, 
females or immature males, showing different: stages in the 
development of the process in the seventh segment of the 
peraeon. 

This species was redescribed and well figured by Mr. W. 
H. Baker in 1910. He states that it is one of the commonest 
marine Isopods of the shores of South Australia. 


PORCELLIO LAEVIS, Latreille. 

ie laevis, Chilton, 1905, Ann. Mag. Nat. Hist., ser. 7, 
vol. 16, p. 

ae obtusifr ons, Haswell, Cat. Austr. Crust., p. 284. 

Two specimens, taken on the shore of Streaky Bay. This 
is an introduced species which is now almost cosmopolitan. I 
have given some notes on its distribution in Australia in the 
paper quoted above. 


METOPONORTHUS PRUINOSUS fprandty, 
Metoponorthus pruinosus, Chilton, 1905, l.c.; p. 431. 
One specimen, on the shore of Streaky Bay. This is 
another introduced species that is now very widely distributed. 


¢ 


38 


Synonyms and notes on its distribution are given in the paper 
quoted. 
List oF REFERENCES. 
Baker, W. H. 
1908—*“Notes on some Species of the L[sopod Family 
Sphaeromidae from the South Australian Coast, 
Part I.’’ Trans. Roy. Soc. S. Austr., vol. xxxii., 
pp. 138-162, pls. in. to x. 
Barnard, K. H. 
1916—‘‘Contributions to the Crustacean Fauna of South 
Africa. 5. Amphipoda.’’? Ann. South African 
Mus., vol. 15, pp. 105-301, pls. 26-28. 
Chilton, C. 
1885—‘“‘Notes on a few Australian Edriophthalmata.”’ 
Proc. Linn. Soc. N.S. Wales, vol. 9, pp. 1035-1044, 
pls. 46 and 47. 
1890—‘‘Revision of the New Zealand Idoteidae.” Trans. 
N.Z. Institute, vol. 22, pp. 189-204. 
1912—‘‘The Amphipoda of the Scottish National Antarctic 
Expedition.’ Trans. Roy. Soc. Edin., vol. 48, 
pp. 455-519, pls. 1. and ii. 
1915—‘‘Deto, a Subantarctic Genus of Terrestrial Isopoda.”’ 
Jour. Linn. Soc., Zool., vol. 32, pp. 435-456, pls. 


39 and 40. 
1917—‘‘Notes on Australian Isopoda: (b) on Deto 
marima.’’ Trans. Roy. Soc. S. Austr., vol. xli., 


pp. 399-404, with text figures. 
1921—Report on the Amphipoda obtained by the F.I.8S. 
“Endeavour” in Australian Seas. Fisheries: 
Biological Results, etc., vol. v., part 2. 
Collinge, W. E. 
1918—On the Oral Appendages of certain Species of Marine 
Isopoda. Jour. Linn. Soc., Zool., vol. 34, pp. 
66-92, pls. 7- 9. 
Haswell, W. A. 
1879—On Australian Amphipoda. Proc. Linn. Soc. N.S. 
Wales, vol. iv., pp. 245-279, pls. 7-12. 
Miers, E. J. 
1881—Revision of the Idoteidae. Jour. Linn. Soc., Zool., 
vol. 16, pp. 1-88. 
Stebbing, T. R. R. 
1900—South African Crustacea, Part I. 
1906—Amphipoda. 1. Gammaridea. Das Tierreich, 21 
Lieferung. 
1910—Amphipoda- of the ‘‘Thetis’” Expedition. Mem. 
Austr. Mus., No. IV., pp. 567-658, pls. 47*-60*. 


39 


THE EXTERNAL CHARACTERS OF POUCH EMBRYOS OF 
MARSUPIALS. 


NO. 3-/SOODON BARROWENSIS. 


By £. Woop’ Jones,*D.Sc.,. F.Z.8., 
Professor of Anatomy in the University of Adelaide. 


[Read April 13, 1922.] 


Of this bandicoot I have so far obtained but three 
embryonic stages for description. For all these specimens I 
am indebted to the authorities of the Perth Museum. As 
opportunities for obtaining further material may be long 
delayed, and as the three stages (17 mm., 77 mm., 92 mm.) 
examined are representative of a long cycle of pouch life, it 
has been thought worth while to record such details as are 
ascertainable from the study of these specimens. 

Har.—Hair is evidently late in development, there being 
no appearance of general body hair at the 77 mm. stage. In 
the embryo of 92 mm. the general body hair is developed, 
and is of the characteristic hispid type and bright tan in 
colour. : 

Har Tracts.—In general disposition the stiff harsh hair 
of the 77 mm. embryo exhibits the utmost simplicity. 

With the exception of one field, the whole of the hair of the 

head, body, and tail slopes uniformly backwards (see fig. 1). 
The exceptional area, which may be defined as the gular field, is 
situated beneath the throat, extending from the angle of the 
mouth to the root of the neck. In this field the hair trend is 
completely reversed. The anterior convergent region beneath 
the chin is marked by the interramal papilla and its vibrisca ; 
the posterior divergent region is situated at the posterior 
extremity of the base of the skull. The lateral margins of 
the area are very definite, and they extend backwards from 
the angle of the mouth practically along the lines of the rami 
of the mandibles (see fig. 2). 
* Upon the hmbs the flow is distal and towards the post- 
axial margin, but in the case of the fore limb a reversal takes 
place at the post-axial margin between the wrist and the 
elbow. Hair is continued to the base of the ungual phalanx 
of both fingers and toes (see fig. 3). 

The sole of the foot is hairy and the arrangement of the 
hair tracts is very definite. A central divergent area is pre- 
sent upon the sole opposite the first digit. Behind this point 
the hair is arranged in two streams running backwards to 
the heel and towards the mid line. This backwardly-directed 


40 
stream meets around the heel with the descending stream from 
the leg. In front of the central point the streams are 
directed forwards along the syndactylous digits, and the fifth 

digit, respectively (see fig. 11). 
The hair colour of the specimen in which hair is 
uniformly developed (Perth, B, 92 mm.) is a bright tan, the 
dorsal surface being of-a darker tint than the ventral surface. 


— 


SUNOA TAL! 
py ivy 
MUNG / 
Wy 
Fig. 1. Fig. 2. 
Hair tracts of the head (from Gular hair tracts (from ~ 
Specimen Male B, Perth). Specimen Male B, Perth). 


CUTANEOUS PAPILLAE AND VIBRISCAE. 


Facial Vibriscae.—The sensory vibriscae and papillae are 
not very conspicuous. By far the largest papilla is the genal 
which gives origin to some six backwardly-directed vibriscae. 
The mystical set is arranged in five rows, of which a single 
papilla constitutes the upper row. The vibriscae are fine and 


‘(qpiog ‘ ee, Uewtoedg utoay) Apog oyy Jo Souq ICE eG Oly 


42 


pale in colour. A single vibrisca springs from the interramal 
papilla. The submental vibriscae are short and insignificant. 
The supra orbital papilla gives rise to two backwardly-directed 
tactile hairs (see fig. 4). 


Fig. 4. 
Facial vibriscae (from Specimen 
Male A, Perth). 


Brachial Vibriscae.—The ulnar carpal papilla gives 
origin to a single very elongated bristle as well as to an 
unusually short one. A_ single well-developed anconeai 
vibrisca is present (see fig. 5). 


Brachial vibriscae (from Specimen Male A, Perth). 


There are no crural vibriscae or papillae present on any 
of the embryos that I have examined. There are no specialized 
cloacal vibriscae. 

Rhinarium.—The rhinarium is naked and elongated. The 
middle line sulcus of the upper lip is well marked and grooves 
the rhinarium to the posterior extremity of its dorsal surface. 
The naked surface is flesh coloured, and it is granulated in a 


43 


regular manner suggesting mosaic. The slit-like narial 
apertures are directed laterally, and their margins are 
entirely naked. The posterior limit of the rhinarium becomes 
more defined in later embryos as the snout region becomes 
pubescent (see fig. 6). 


Rhinarium (from Specimen 
Male A, Perth). 


External EFar.—In no embryo of the genus /soodon that 
I have so far had the opportunity of examining is the ear 
laid forward at the younger stages. In J. barrowensis the 
pointed ear stands well out from the side of the head with the 


Fig. 7. 
The form of the external ear 
(Embryo Male A, Perth). 


tip directed backwards. The processus antihelicis (the so- 
called metatragus) is large in all stages; the characteristic 
adult twist in its length becoming more pronounced as growth 


ad 


proceeds. Nearer to the external auditory meatus than the 
metatragus of the taxonomist is a second and smaller process 
of the antihelix which, by becoming separated from the main 
process with the enlargement of the auricle, leads, in the 
adult, to the formation of a pit. between the two processes. 
The ‘‘deep hollow” described in this area of the adult ear 
is, however, a secondary feature formed by the relative growth 
of the surrounding parts. , ‘ 
Two genuine pockets are, however, present in the auricle. 
The first is the common mammalian pocket in the posterior 
portion of the helical margin, and which is generally known 
as the sulcus auris posterior. The second pocket is a remark- 
able one (marked A in figs. 7 and 8) in the centre of the 
developing tragus. This tragus pocket becomes covered by the 
hair of the cheeks in the adult, nevertheless it remains a 
permanent and remarkable feature of the external ear. 
Manus.—The digital formula is 2=3>4>5>1. Claws 
are developed at the 17 mm. stage upon digits 2, 3, and 4; 
but 1 and 5 are clawless. The digits are fusiform, tapering 
towards their distal extremities; there are no definitely 
developed apical pads. Three basal pads are developed, one 


Fig a8: 
Form of the external ear (from 
Specimen Male B, Perth). 


being opposite the base of each clawed digit. The skin of the 
palm is granular (see fig. 9). — 

Pes.—The digital formula is ASS HS 20> 1. 

The outstanding features of the foot are the great size 
of the fourth digit and the reduced condition of the first, 
which bears no claw. The digits are fusiform. Three basal 
pads are present, one being at the base of digit 5, a larger one 
at the base of 4, and a small one at the base of the syndactylous 
elements. 


‘ws 


<ssiaidl Pon 


45 


In the 77 mm. embryo the sole is uniformly granular, 
but at 92 mm. hair has clothed a large area of the plantar 
surface of the pes and of digits 2 and 3 (see figs. 10 and 11). 


SE 


f 

j 
i, 

y 


Yow 
\ 
JEN \\| 


G 


\ 


is 


Fig. 9. NY 
Palmar surface of the Fig. 10. | “Wig. HU. 
left manus (from Plantar surface of the Plantar surface of the 
Specimen Male A, left pes (from Speci- ~ left pes (from Speci- 
Perth). men Male A, Perth). men Male B, Perth). 


External genitalia.—The genital tubercle of the male is 
extra-cloacal at the 17 mm. stage, and intra-cloacal at the 
77 mm. stage. The opening of the pouch in the female 1s 
directed downwards and backwards. 


46 


A GEOLOGICAL TRAVERSE OF THE FLINDERS RANGE 
FROM THE PARACHILNA GORGE TO THE LAKE 
FROME PLAINS. 


By Proressor Wa.tTEeR Howcain, F.G.S. 


[Read May 11, 1922.] 


PRAT y: 
CONTENTS. 

PAGE 
I. Parachilna Gorge ... he oe x ers eae 
II. Horne’s Camp vy ae es 
Ili. Blinman and Nena choad oe “al here n o.: 
IV. Blinman to Reap-hook Range oe wie Je uate 
V. Trip to Five Miles North of Blinman _... ao) 
VI. Trip to Patawarta Hill a a ise as 
VII. Western Side of Blinman ... : vac eee 
VIII. South Blinman, and Road to Wirrealpa ak, [Oe 
IX. Wirrealpa and Neighbourhood _... ey oe OD 
X. Up a Tributary of Wirrealpa Creek ... 66 

XI. The Obolella Limestone on the Road to the Old 
Wirrealpa Station 5 67 
XII. The Old Wirrealpa ations ee me eee 
XIII. Visit to Mount Chamb>-rs Creek ae <. , 
XIV. Mount Lyall ... 70 

XV. From Wirrealpa to the “Big Hil? on the Ras 
to Blinman 72 

XVI. Visit to the Gemdeenne TRemeas. Balcoraca 
Creek, and the Wilkiwillina — et See ene 
XVII. Lithologic Features ... ae ee 
XVIII. Tectonic Phenomena hove an ae Fe ae 


In 1906 the present writer crossed the Flinders Range 
from Parachilna, westerly, to the eastern slopes border- 
ing on the Lake Frome plains. The journey was done mostly 
on foot. Publication of results was deferred with the hope 
that opportunities might arise by which the geology of the 
country could be still further investigated and descriptions 
made more complete. As this is now unlikely, the present 
notes are placed before the Society as a summary of the work 
done at the date mentioned, incidental to defects which must 
necessarily accompany observations made on a single traverse 
of the region. 


(1)Some preliminary notes of this journey were included in a 
paper by the author, read at the Adelaide meeting of the Aus. 
Assoc. for Adv. of Science (1907), on ‘‘A General Description of 
the Cambrian Series of South Australia,’ pp. 414-422. 


47 


The typical structure exhibited by the Flinders Ranges 
takes the form of broad anticlinal and synclinal folds which, 
by complicated directions of pressure, frequently produce peri- 
clinal domes with complementary saucer-shaped depressions, 
the latter being locally known as ‘‘pounds.’”’ The geological 
section, now under description, is transverse to one of the 
most extensive dome-structures in the ranges, the centre of 
the dome being situated, approximately, near the township of 
Blinman, with the superior beds dipping away in circles around 
this centre. A few miles to the south of Blinman is the 
Wilpena Pound, formed by a complete circle of mountains 
with the gap made by the Wilpena Creek, the only means 
of ingress and outlet to the basin. At Mernmerna, on the 
great northern line, the hills on the eastward side of the 
line form very steep escarpments with rugged peaks, forming 
the western limits of the Elder and Wilpena Pound Ranges. 
This precipitous face continues, northwards, to Parachilna, 
as a fault-scarp, making the eastern boundary of the great 
rift valley of South Australia in that direction. 

As the present paper is based on a single visit to the 
locality, and an interval of about sixteen years has passed 
since the observations were made, the paper is practically 
limited to the itinerancy and the field notes made at the time. 
The Geological Section, published herewith, was drawn soon 
after the author’s return to Adelaide. The newer, southerly 
road was followed going out, and the older, northerly road 
on returning, when the journey was made by coach. 


I. PARACHILNA GORGE. 
ENTRANCE TO THE GORGE. 


The Parachilna railway station is situated on the plains 
skirting the eastern side of Lake Torrens, about seven miles 
from the foot of the ranges. The gap in the ranges, east of 
the railway station, has been cut by the Parachilna Creek, 
forming a narrow and very picturesque gorge. In approaching 
the gorge the first rocks met with are limestones that outcrop 
on the road, near an old house in ruins. These are sub- 
crystalline, of a coarse-marble kind, much broken and 
penetrated by veins (dip S. 20° W. at 46°), underlying which 
are limestones containing Archaeocyathinae. The country: 
along the face of the great escarpment is much faulted. 
Following the western escarpment, going south, in a second 
spur, the fossils gradually disappear in a dolomitic matrix, 
the fossils occurring in every stage of alteration as they become 
absorbed into the matrix. [On this spur is an isolated group 
of sandstone boulders, some of which are of great size.]| In 
following the line of outcrop, southwards, there is a narrow 


48 


belt of dark-coloured oolitic and laminated limestone, 
apparently brought in, as a repetition of the strata, by a 
strike fault. The limestones can also be traced up to the 
great pinnacle of quartzite which cuts them off on the southern 
side, by a dip fault, the quartzite having an apparent dip 
of 85°, westerly, on its abrupt face to the west. The outlier 
of oolitic and laminated limestones, brought in by the strike 
fault;(has.a dip.S.. 30° W «at, o0e. gots NM. 
The junction of limestone against §.E, 

the quartzite on the fault plane, is 
a rotten brecciated rock. The last 
solid rock on the limestone side, is a 
marble (see fig. 1). 


THE GORGE. 
On entering the gorge the lime- 
stones are seen to outcrop on both 


sides of the creek, mostly on the 4 AV) 


southern side, where they form a 
ridge about 200 ft. high and make a 
spur, running westward, to the 
plains, where they pass from view 
under alluvium and sand. The lime- 
stone facing the plains has a strike Outlier by complex fault- 
BE. 25° S., dip 45° westerly. The i2g of limestones pinched 
gorge road intersects the limestones ‘” betes eae 
obliquely to the strike of the beds and supplies an interesting 
section, as detailed below : — 

(a) The bottom series in the limestone belt begins about 
half a mile from the entrance of the gorge, resting on thick 
quartzites which rise to a great height: dip S. 30° W. at 60°. 
The beds are characterized by a series of dark-coloured oolitic 
limestones, separated by earthy bands, or beds, which continue 
as a cliff facing the creek for a distance of 300 yards: dip 
60°-75°. 

(6) Blue and buff limestones, dolomitic; nodular, 
stalactitic, laminated; more rarely, finely oolitic in struc- 
ture; white crystalline limestone, passing into white to brown 
dolomitic marbles, with reduced angle of dip. About 300 
yards of outcrop. 

A small tributary creek dissects the cliff at this point, 
making a gap 36 yards wide, but the limestones continue 
up this creek. 

(c) Pink and yellow marbles continue for a distance of 
about 50 yards, passing up into very solid and continuous 
white to buff marbles. Width, 90 yards. 

Another wash-out by a small tributary creek, 75 yards 
wide, with limestones passing up the creek. 


49 


Granular marble shows again in cliffs on opposite side 
of wash-out. Dip S.W. at 40°. These beds continue for 
180 yards to another small affluent, where the Archaeocya- 
thinae beds make their appearance with a strike EK. 20° S. 


(d) Archaeocyathinae limestones, showing extensive 
development near the outlet of the gorge. The base of these 
beds cannot well be determined as the fossils gradually dis- 
appear, losing their organic structures by conversion into 
marble or dolomite. This occurs both at the base of the 
fossiliferous beds as well as at their upper limits.. 


Immediately on the eastern side of the great limestone 
series, the Parachilna Creek has broken through a great wall 
of quartzite, which towers to a great height on either side. 
There follow, in descending order, thick shales with some flags 
(dip 70°-80°) up to the prominent and peaked hill of quartzite, 
near the reservoir; underlying which are shales having a 
strike almost parallel with the road, and a dip of 90°. These 
shales are often sharply curved and continue in the section 
to the mouth of the Oratunga Creek, having a dip of from 
80° to 90°. From this point, by a sharp bend in the road, 
the latter follows the strike of the beds, which there have a 
dip of 75°. The road then crosses the creek and takes a 
sharp curve round a spur of quartzite, 50 ft. in thickness, 
with a dip from 70° to 80°. In this angle there is a fault 
associated with strong V-shaped contortions, the beds being 
shales with hard, thin quartzites having a dip of 45°. Ata 
short distance from the preceding the road crosses the creek, 
-a second time, where shales have a dip of 60°. At the third, 
and last time that the road crosses the creek, the shales have 
a dip of 80° to 85°, with a wavy structure. 


At a distance of three miles from the mouth of the gorge 
limestones once more begin to show themselves in the section. 
At a sharp angle of the road, just past Mount Mary, there 
are beds of pink-coloured limestone seen on the road. Shales, 
with a dip of 45°, occur for a distance of three-quarters of a 
mile to the ‘‘Dairy,’’ where the road is close to the Parachilna 
Creek and is at the foot of the ‘‘Big Hill.’’® The dip 
decreases towards the ‘“‘Dairy.”’ 


Calcareous grits and arenaceous (oolitic) limestones occur 
very commonly on the western side of the ‘‘Big Hill.’’ There 
also occur thick flaggy quartzites, 200 ft. in thickness, over- 
lain by pure limestone and gritty oolitic limestone: dip S.W. 
at 25°. These gritty limestones have a great development 
on the creeks which pass between the road and the Parachilna 


(2) This is quite distinct from a hill of the same name situated 
between Blinman and Wirrealpa. 


50 


Creek, and these, interbedded with flags and quartzites, 
represent a series some hundreds of feet in . thickness. 

Leaving the ‘‘Dairy,’’ going east, the road follows the 
creek, taking the rise of the “‘Big Hill,’”’ showing, in suc- 
cession, for about a mile, shales (dip 45°); blue lmestone, 
seen in creek; shales (dip 15°-20°); lhmestone (dip 15°); 
calcareous grits (dip 15°-20°). The road reaches its greatest 
elevation between the Gorge and Blinman at the north turn, 
or bend, known as the “Big Hill,” or the ‘‘Seven-mile Hill” 
(measured from Blinman). At this point there is a reef of 
iron and lime carbonates crossing the road. The Seven-mile 
Hill is chiefly marked by quartzites, 100 ft. in thickness, 
with a dip S.W. at from 10° to 15°. [Top-bed (?) calcium- 
carbonate.| On the eastern side of the ‘‘Big Hill” is the 
“Snake Bend,’ and at 6? miles from Blinman, an arenaceous 
limestone, 5 ft. 6 in. in thickness, interbedded with flaggy 
quartzites, occurs. At 6 miles from Blinman, a small quarry 
is worked in a siliceous limestone, by the side of the road: 
dip S.W. at 35°. The stone is used for road metal. 


Il. Horne’s Camp. 


On the evening of the first day out I reached a spot 
known as “Horne’s Camp,’’ situated on a creek, tributary to 
Parachilna Creek,‘5) that crosses the road about 4 miles to 
the westward of Blinman. A party of Government ‘‘road- 
men’’ was camped on the creek, with whom I found accom- 
modation for the night, and spent the following day examining 
the creeks in the locality. 

Facing the camp, from the southward, is a long escarp- 
ment that follows the strike and has the appearance of a great 
rampart giving a clear exposure of the beds. The latter 
consist of argillaceous and arenaceous flags that split per- 
fectly on the bedding planes. They can also be studied in 
the creek banks, near the camp. Here bluish quartzites with 
partings of softer material outcrop with a dip S., and 
S. 20° E. at 30°. Followed the creek, downwards. At 
about 400 yards from the camp, a 4-ft. banded limestone 
occurs that splits up easily into flags. At two-thirds of a 
mile, in the same direction, dark-coloured fissile flags occur, 
much like Willunga ‘“‘slate,” having a dip S.W. at 20°. 
Here the stream that I was following joined a larger creek 
which came in from the east. At three-quarters of a mile 
from the above junction, going down stream, a quartzite is 
met with, 50 ft. in thickness,” passing in its upper portions 


(3) The Pastoral Map on which is shown the two creeks referred 
to is entirely unreliable as to their respective directions. 


51 


into flags with a wavy structure, slightly false-bedded. The 
dip varies from 17° to 20°. The beds that follow are a 
lenticular dolomite, in quartzite, situated near sharp bend 
of the stream (seen on the rise); calcareous bands, in flag- 
“stones, showing lamination by weathering; the flagstones, in 
ascending series, become more calcareous and pass into 
calcareous flagstones. A waterfall, having a height of about 
20 ft., occurs on this creek, about one-eighth of a mile before 
‘its junction with Parachilna Creek. The section continues 
in flagstones, quartzites, gritty limestones, and calcareous 
grits: dip S.W. at 20°. Some of the gritty beds are from 
10 ft. to 15 ft. in thickness, and, in places, show ripple 
marks. The hill, adjacent to waterfall, consists mainly of 
gritty limestones, and has a height of from 200 ft. to 300 ft. 

In the main creek, around a westerly and southerly bend, 
there is a sudden change in the dip, passing into intense 
folding and a throw down, by fault, at 90°. The fault area 
is well defined by walls that are vertical and 14 ft. apart— 
the fault area is filled by fault-rock. On the eastern side of 
the fault the dip is N. 65° W. at 40°. On the western 
side of fault the dip is S.W. with numerous small thrust folds, 
which extend for 100 yards. At the next bend, one-eighth 
of a mile below the previous observation, the dip is W. at 
30°. At one-eighth of a mile, further down, rapids are 
formed by a bar ofgritty lmestone, overlain by flags, and 
include a thin bed of volitic marble, 1 ft. in thickness, and 
having a dip W. at 30°. At another one-eighth of a mile 
distance, a second strong bar of gritty limestone occurs, 
making a cliff-bank 500 ft. in height: dip S. 60° W. at 30°. 
The beds form a synclinal fold. The beds underlying the 
synclinal curve are oolitic limestones, 20 ft. in thickness, 
and inferior to these there are impure wavy limestones, with 
a dip S.W. at 20°. 

At this point I left the main creek and followed up a 
small tributary which drains in from the north-east. In this 
_ ereek a striking fold occurs which crosses the stream, on the 
eastern side, the dip is W. at 14°; and on the western side, 
the dip is N.W. at 80°, in flagstones. Higher up the creek, 
there is another throw down to N.W. at 67°-80°; the section 
showing dolomitic limestone in beds from 12 ft. to 18 ft. in 
thickness. At the head of the creek there are thick quartzites 
which rise to a crest of about 300 ft. in height, with a 
shoulder of quartzite, at a lower level; and a yet lower one, 
of flagstones, which, latter, come down to the level of the 
road about half a mile westward of the camp. 

Spent the second night in camp with the road-men. Next 
day cut across country to Blinman. Examined travertine 


52 


near camp. It is exposed in cliffs of the creek both above 
and below where the road crosses the creek. The beds are 
horizontal and up to 12 ft. in thickness; the base of the 
beds is, usually, a few feet higher than the normal level 
of the creek. The limestone varies from soft, loosely-cemented- 
globular concretions to a compact rock, often finely brecciated. 

Went up small creek, adjacent to the camp, going north. 
At. a distance of 200 vards, up the creek, the rocks were found 
to be argillaceous flags: dip S. 20° W. at 75°, increasing to 
85°. At a further distance of 200 yards, further up, thick 
beds of dolomitic limestone, in rolling folds, with an average 
dip of 40°. These beds show some extraordinary effects of 
crush—laminated, contorted, broken, passing into crush 
conglomerate in which dolomitic limestones and shales are 
mixed together. This broken area extends for a width of 
50 yards, giving no evidence of dip, and is underlain by con- 
torted slaty flags, with a‘dip S. 20 E. at 80°. 

A little higher up the creek, another bed of dolomite 
(or dolomitic limestone), 9 ft. in thickness, is included in 
disturbed and broken slates which are in vertical position. 
Thick quartzites follow a ridge that forms the crest of a 
very pointed and conspicuous hill on the north side of the 
camp, and are underlain by very thick dolomite, with a dip 
of 25°. These beds occupy the creek for one-eighth of a 
mile, are finely crystalline in texture, and, in quantity, 
sufficient to rebuild the Westminster Houses of Parliament. 
The thick dolomite is followed, in descending order, by 
laminated shales, at a dip of 80°, including a bar of dolomite, 
and these are underlain by a dark-coloured contact (garnet) 
rock, which has undergone alteration by contact with an 
igneous dyke. | 

A greenish, basic dyke, 24 yards wide, runs up the face 
of the hill on the eastern side of the creek, and outcrops on 
top, on the south-western side of the saddle from which rises 
the precipitous peak of quartzite, already referred to. It is 
20 yards wide at the top of the hill and throws out lateral 
dykes. 
ts Higher up the creek the section shows rotten purple 
shales having a strike S. 20° W. with dip at 90°. 

The country now becomes more or less reticulated with 
basic dykes, over a breadth of a quarter of a mile, or more. 
One very prominent dyke that intersects the creek is 25 yards 
wide, bordered by shales on the one side and quartzites on 
the other, which show contact metamorphism. 

Followed up the main north-eastern creek for a while. 
In the alluvial of this and other creeks were pebbles of 
brecciated limestones, as well as ‘“‘greenstones’’ derived from 
intrusive dykes. Crossing the low range, on the eastern side, 


53 


basic dykes outcrop with a strike east and west. Further 
intrusive dykes were seen on the next range, 2 miles distant 
from the most westerly outcrops that were noted. Similar 
intrusions were observed crossing the old Blinman road in 
several places, in one case giving a width of 60 yards. The 
sedimentary rocks met with in this cross-country journey to 
Blinman were some prominent outcrops of quartzite, flaggy 
sediments, and small dolomitic limestones. 


TII. BuinmaN AND NEIGHBOURHOOD. 


The Blinman township, according to official figures, -is 
situated 2,020 ft. above sea level and 1,555 ft, above the Para- 
chilna plain. The mine is in disturbed country, and the 
copper ores occur mostly in a dolomitic limestone near its junc- 
tion with flaggy slates. These features can be well seen in the 
open cut where the limestone makes the foot wall and the slates 
the hanging wall. The dip varies from 65° to 75°. The cap- 
tain in charge stated that the shaft cut the limestone at a 
depth of 50 fathoms and passed diagonally through it to the 
70-fathom level. The ore in the upper part is in the form of 
copper carbonates and black oxide, which intersect the lime- 
stone by reticulation of large and small veins. The average 
width of the payable cupriferous zone is 14 ft. The ore 
sometimes lies in flat shoots, the thickest part of the shoot 
being from 1 to 2 in. The ore body seems to be limited by 
a eross-fault with an east and west strike on the southern 
side of the mine. From the nature of the ore distribution 
the whole of the mineralized country is worked as stock- 
works and smelted. As the ore is carried in limestone, ores 
of a siliceous nature are bought, when possible, for fluxing 
purposes, and sandy shales are also quarried and used for a 
similar purpose. [Since -my visit the mine has been, unfor- 
tunately, closed. | 

The hills on the eastern side of the Blinman Mine consist 
of shaly flags and crushed dolomitic rock. One hill, just 
east of the mine, exhibits a small syncline on its summit, best 
seen from the southern side: dip N. 65° E. at 75°, and S. 
20° E. at 60°. Half a mile to the northward of the mine is 
an igneous dyke, 18 yards wide, with a strike 20° S. of W. 
On the north side of the dyke there are strong flaggy quartzites 
that make a prominent hill and carry a thick band of lamellar 
hematite. A similar quartzite follows around the western 
side of the mine, and at a distance of one-eighth of a mile 
from the mine, in the same direction, there are strong beds 
of dolomitic lmestone, much crushed, in association with 
igneous intrusions, of which there are three circular bosses 
(? chimneys), forming, by position, a triangle, about 100 


54 


yards distant from each other, each having a re 
of about 50 yards. 

Crossed country to creek, on road to Parachilna, one mile 
east of Horne’s Camp, and followed the road and creek back 
to Blinman. Laminated shales are seen in creek, on southern 
side of road, with a dip 8.W. at 15°; which suddenly increases, 
by a twist, to 73°; and, further up, to 85°; then swings 
around to E. .20° S. at 50°. Where the creek crosses the 
road the shales are intensely broken and reunited in large 
angular pieces. Following which, where the creek forms a 
loop on the southern side of the road, the shales, which are 
at first much shattered, pass into a synclinal fold with a 
high dip, and then to a lower angle with dip S. A suc- 
cession of basic igneous dykes formed the chief feature for 
some distance on the same line of section. /urst dyke occurs 
at bend of the road in creek, on northern side, and is 40 yards 
wide with a dip of 85° W. The hanging wall to the dyke, 
on the western side, consists of laminated shales, that are not 
disturbed, having a dip S.W. at 45°. The dyke is under- 
lain, on the eastern side, by shales which are greatly crushed 
and broken up. This broken rock shows an outcrop, up- 
creek, of 45 yards; then follow laminated, decomposed, yellow 
shales, with dip N.W. at 80°; underlain by thinnish dolomitic 
rock, which is broken. Second dyke, situated 200 yards 
higher up than the first dyke (14 miles from Horne’s Camp). 
It is fine-grained and very basaltic like, is 60 yards wide, 
and a strike E. It is bordered by shales that are brecciated, 
with dip S. 20° W. at 83°. A little further on the road, a 
strong outcrop of quartzite is seen to come down the hill face 
on the western side of the road, but is cut off at the creek, 
and in its place, on the eastern side of the creek, is a fault 
rock, much broken for 30 yards, followed by dolomitic 
rock, with dip W. 20° N. at 75°. This, again, is followed 
by laminated quartzites, with dip N.E. at 60°; then swings 
around to N. Third dyke, situated about 2 miles from Horne’s 
Camp, is 42 yards wide, with a strike W. 20° N., follows 
a small creek, and can be traced across larger creek, into 
hill, on the western side. Fowrth dyke, situated about 200 
yards beyond the third dyke, is 5 yards wide. There is a 
great show of crush rock on hill, on the eastern side, down 
to the road, consisting of shales and dolomitic rock in a 
mixed condition. For the next 200 yards on the road, shales 
and some dolomitic limestones occur. Fifth dyke, 24 miles 
from Horne’s Camp 120 yards wide. Junction of rock on 
southern side gives dip E. 20° N. at 65°. Two hundred 
yards further, blue limestone is seen on road, dip S.E. at 50°, 
with crushed dolomitic rock on top. Quartzites in shales, 


55 


- 


with dip N. at 45°, make a low rise in the ground. One and 
a quarter miles from Blinman there is a high and bold ridge 
of slaty dolomitic rock, somewhat broken in places, underlain 
by shales: dip S.W. at 80°. Last prominent ridge, on the 
eastern side of road, in a direction N. 20° E., consists of 
laminated shales, thin quartzites, and dolomitic rock in places. 
At one mile from Blinman are shales, dip 8.W., with broken 
beds of slaty dolomitic rock. From this point the ground is 
low, undulating, and grassy. 


ITV. BLINMAN TO REAP-HOOK RANGE. 


Trip southward (4 miles) to Reap-hook Range. So called 
from its resemblance to the tool—a handle, and great curve 
for blade. Also known as Patterton Hill and Mount Emily. 
Drove out to Patterton Spring and then went one mile across 
to Reap-hook Range. The latter has a very striking rock 
face 500 ft. in height. The top beds consist of 25 ft. of 
impure arenaceous limestone having a vertical scarp: dip 
E. 25° 8. at 5°. Beneath which is a steep scarp of purple 
shales and laminated flaggy shales. The latter are also seen 
at the Patterton Spring (mentioned above), where they dip 
S. 20° E. at 10°. In retracing my steps, on foot, from the 
Springs to Blinman, underlying the slates, just mentioned, is 
a dolomitic rock, then a limestone which weathers with a 
dark-coloured smooth surface, similar to the Archaeocyathinae 
limestone, but has arenaceous lines in relief that follow, 
generally, a circular outline; then an arenaceous limestone, 
showing an outcrop of about 100 yards, with dip at a low 
angle. A thin quartzite occurs in the limestone series, which 
is underlain, again, by thick arenaceous limestones; then 
ochreous limestone with vein of siderite. For the next half- 
mile there were noted calcareous beds, separated by thin 
beds of shale; then quartzites and calcareous grits: dip S.E. 
at 12°. 

There follows a relatively flat country, in which shales 
are first met with, then solid limestone showing wavy struc- 
ture and is sometimes arenaceous, which continues to small 
creek, three-quarters of a mile before reaching Youangera 
Spring. On the Blinman side of the creek another strong 
limestone is seen on a prominent rise, just before sharp bend 
in the road (between Sections 64 and 69), with shale on top 
and is underlain by quartzite, with a dip S.E. These beds 
are followed by a somewhat lower hill, consisting of flaggy 
shales; and then, strong calcareous grits passing into brecci- 
ated limestone, which latter makes a bold ridge that crosses 
a creek that is tributary to the Blinman Creek, about a 
quarter of a mile above the road crossing. Some fine springs 
occur in the creek a few yards above the crossing. 


56 

Soft shales are on the flat skirting the eastern side or — 
the limestone range, just mentioned (dip E. 20° N.), and 
these are faulted against shales with thin beds of ferruginous 
limestones which dip 8. 20° W. Both these sets of outcrops 
are at high angles, as seen on the flat, and are also much 
curved. 

At the road crossing the tributary creek, near the 
Youangera Spring, flaggy shales dip E. 20° S., at from 40° 
to 45°. Here a very curious white limestone is seen resting 
unconformably on the shales. The limestone rolls a little, 
with anticlinal and synclinal curves, and is eroded where the 
curve passes above the normal level of the ground. It is 
compact, somewhat nodular, .and includes numerous frag- 
ments of shale. It is veined with crystalline matter and has 
manganese oxide stains. The question as to its origin carries 
some doubt, but it is most probably a travertine limestone 


with certain unusual features. It is seen on the north side . 


of the creek, and can be traced to the junction of the two 
creeks, a distance of about 100 yards, beyond which I did 
not continue my observations. The bed does not seem to 
rest on calcareous rocks; it is from 6 to 8 ft. in thickness— 
limestones occur on the scarp face about one-eighth of a mile 
to the eastward. It seems probable that it is a travertine 
deposit laid down, at a somewhat distant period, by spring 
waters fed from the calcareous beds of the scarp that exists 
on the eastern side, perhaps before the scarp had retreated 
as far as at present. The creek that has cut its way through 
this peculiar limestone gives no evidence of carrying any 
quantity of calcium carbonate in solution at the present time. 
On the western side of the crossing, in the same creek, there 
are similar laminated shales as occur above the crossing and 
are dolomitic, in places: dip E. at 40°. There is also an 
overlying limestone, on this side, but it is not so developed 
or so compact as in the higher position in the creek, described 
above. 

The road, after crossing the creek, has a trend more to 
the west and passes over a ridge of shales, sometimes cal- 
careous, which pass under the limestones of the scarp, described 
above: dip S.E. at 50°. Beyond the last-named ridge, a 
12-ft. bed of limestone occurs in the shales, followed by flaggy 
quartzites, which make a bold hill on the western side of the 
road: dip E. 20° S. at 75°, increasing to 90°. On the other 
side of the range—in lower ground—the rocks are somewhat 
broken and have an easterly dip. 

Passing into the valley of the Blinman Creek, shales and 
flaggy quartzites form the outcrops, the quartzites carrying 
the fine dark lines similar to those seen in the quartzites on 


Don 


57 


hill west of South Blinman, and also west of the Blinman 
Mine. 
THE BLINMAN CREEK. 


The Blinman Creek, near the townships of North and 
South Blinman, supplies exposures illustrating crush-rock to 
an extreme degree. The purple shales, particularly, are con- 


verted into crush-breccias and crush-conglomerates in which 


the original bedding is entirely obliterated, or is present only 
in isolated fragments. The locality is also greatly inter- 
sected by intrusive dykes. | 

Going south from South Blinman (in creek) the purple 
shales are overlain by flaggy shales: dip W. 10°:S. at 80°. 

(1) A great basic dyke crosses the creek forming a pro- 
minent ridge 30 ft. in height and 100 ft. in width. The 
dyke cuts across cupriferous flaggy shales, the latter having 
a dip of 70° W. of N. at 60°. 

(2) Two hundred yards lower down the creek another 
dyke crosses the stream—on the eastern side, measuring 15 ft. 
in height and 90 ft. in width. On the southern side of the 
dyke purple shales form a cliff face in the creek and are 
intensely broken and brecciated. The dyke crosses to the 
western side, where it is seen on the rise of the hill. The 
strike is N.E., and follows along the slack ground. 

(3) On the western side of the creek, about midway 
between the two dykes just referred to, is a circular outcrop 
of igneous rock, 100 yards in circumference. The stone is 
scoriaceous, in parts; whether the vesicular structure is due 
to gas cavities, or spaces left by the decomposition and removal 
of included crystals, is not quite clear, but the cavities have 
been subsequently filled, in some instances, with Fe CO, and 
other crystals. The circular outline of the outcrop suggests 
the possibility of its being an old volcanic ‘‘neck.’’ 

The prominent hill that is on the western side of South 
Blinman has quartzites at its summit, and, for the most part, 
on its southern face also: dip 10° S. of W. at 75°. The 
quartzite carries dark lines (a common feature in the quartzites 
of the Upper Cambrian series), and there is- also a dolomitic 
limestone, much contorted and irregular, that outcrops on 
the southern and south-eastern flanks of the same hill. The 
dolomitic bed has a rolling strike of N.W. and 8.E. 

A ridge of hills runs parallel with the road between South 
Blinman and North Blinman, on the eastern side, consisting 
of flaggy shales and quarizites, with dip N.E. at 75°. 

’ Following the Blinman Creek to North Blinman, a great 
development of crush-rock occurs both in the creek and on 
the flanks of the low ranges on the western side. The rocks 
consist of siliceous shales and thin dolomitic hmestones, in 

C 


58 


which the ‘strike and dip vary with every few yards, from 
horizontal to vertical, the dip tending east and west. From 
the Government well, situated in the creek, the beds become 
more regular (going north), the siliceous shales showing a 
dip N.E.-at 50°. 

On the bank of the creek are some very large spheroidal 
masses of siliceous quartzites, and these occur again, higher 
up the creek, containing dark lines, and have a dip N. 20° W. 
at 45°. This spheroidal weathering in homogeneous siliceous 
rocks, as well as the dark lines, often cross-bedded, are very 
characteristic features of the Upper Cambrian beds of the 
Flinders Range. 


V: Trre Five Mites Norty or BLinMAN. 


Followed the road on the eastern side of the mine, 
which passes over a flat and trends in a north-westerly direc- 
tion. Dr. Lander drove me to Little Willigon Creek, which 
is separated from the Willigon by a ridge, at about 5 miles 
from Blinman. Left the conveyance and proceeded on foot. 
The Little Willigon Creek cuts through the ridge mentioned, 
in a small gorge, with shales on the one side and limestone 
on the other. The limestone, which makes the ridge, at the 
gorge, shows a structure of concentric lines like globules 
1 in. to 2 in. in diameter, which weather into depressions ; 
it has also inclusions of shale in angular fragments. It is 
underlain by flaggy shales that dip N. at 35°. 

By climbing the ridge between the Little Willigon Creek 
and the main Willigon Creek (a distance of about three- 
quarters of a mile), from its highest point the geological 
structure of the country could be well seen. An imposing 
range on the north side (4 miles distant) marks the limit of 
vision in that direction with a steep scarp face on the southern 
side, apparently composed of flaggy shales with interbedded — 
impure limestones. Then followed an inner range of rounded 
hills covered with green feed, approximately 2 miles wide. 
From the physical features I concluded that this area was 
composed of purple shales. Another range, at a shorter dis- 
tance, occupied the space down to near the Willigon Creek, 
and showed a steep face to the southward composed of flags 
and thin limestones. All these outcrops to the north of the 
creek could be distinctly seen to dip northwards, giving a 
section of 4 miles in diameter. 

On the southern side of Willigon Creek there is a suc- 
cession of hills increasing in altitude towards Blinman, con- 
sisting chiefly of impure limestones with some flaggy shales. 
About one mile from the gorge of the Little Willigon Creek 
the road crosses a small tributary of the latter in which 


59 


limestones make a great development on the northern part of 
the hill, with a dip N. 20° E. at 23°. The limestone has 
a concretionary structure and is very brittle with a spheroidal 
fracture. The limestone is underlain by shaly flags. At a 
quarter of a mile further the road crosses the tributary stream 
again, at a bar formed by another limestone that is impure, 
carrying streaky lines and reticulations of an earthy nature, 
in relief, as well as small stones. This limestone outcrop is 
53 yards wide, with a dip N. 20° E. at 30°. Within a few 
yards it is followed by another hmestone, quite as thick as 
the preceding, and forms a scarp face which runs parallel to 
the stream and road for 14 miles; the road then takes a 
southerly turn. 

On the southern side of the great limestone. series there 
follow, in descending order, a thick series of flaggy shales 
-with ferruginous dolomitic limestones and grits in prominent 
edges ({?]2 miles across the strike). The road rises to a 
high point, where the shales dip N. 20° E. at 30°, and are 
overlain by grits and a ferruginous dolomitic rock. 

Coming down the hill on its eastern side the strike swings 
round a little, with a dip N.E. at 25°, which has the effect 
of bringing the limestone once more across the road, where 
the latter crosses a large creek. The road continues on the 
line of junction between the limestone and shales for over a 
mile. The road crosses the creek for the last..time, where 
the shales have the same dip as in the last reading, viz., 
N.E. at 25°. | 

The road now curves round to tke south towards Blinman. 
Flaggy shales continue on low ground. About one and a half 
miles from the Blinman Mine, situated near the road, in a 
small wash, there are gritty rocks, much broken and twisted, 
in an apparently vertical position, and, mixed with these 
broken beds, is a deposit of small quartz crystals, separately 
developed, making a width of 10 yards, and extends still 
further in patches. 

About a mile from the Blinman Mine, on the western 
side of the road, there is a great spread of ‘eritty limestones 
on the flat, making an outcrop 200 yards in width, underlain 
by flaggy shales, best seen on the rise of the hill, having a 
dip N. 30° E. at 25°. The same shales are seen in the creek 
on the eastern side of the rise, with a dip N-E. at 27°. 

The gritty limestone, just referred to, appears to be cut off 
by a strike fault on the eastern side and is probably a repe- 
tition of the limestone of the range seen to the north. The 
disturbed strata in the valley (referred to above) may be 
regarded as suggestive of such a fault. The associated shales 

“pass up into quartzitic rocks on the rise, with shales on the 

c2 


60 


other side in a much disturbed condition: dip 8. 20° E. at 
from 40° to 50°. 

On the next flat (and, apparently, on the rise to the 
west) there is a thick limestone with concentric structure, 
in an outcrop of about 200 yards. This bed is underlain by 
rotten shales, seen in a small quarry on the top of the rise 
on the road, above the township, with a dip N. at 45°. On 
going down the slope to Blinman there are evidences of 
dolomitic limestones, which probably may be correlated with 
a similar bed at the mine. 


VI. Trip tro PatrawarRta HItu. 


Patawarta, as seen from Blinman, looking northwards, 
has the appearance of a great wedge-shaped pinnacle, rising 
conspicuously above all the surrounding hills, being the 
highest point of a bold range of quartzite having its scarp 
face to the south and dip slope to the north. Mr. J. V. 
Whyte, of Angorigina Station, kindly drove me out a dis- 
tance of 12 miles to visit this interesting hill, which official 
Survey Reports state to be 3,060 ft. in height. 

The road lay through the Nildottie Gap, up the valley 
of the Artimore Creek, past Artimore Head Station, and 
over the shoulder of the Patawarta Hill, on its western side. 
The journey took in country seen to the north on my trip 
to the Willigon Creek. | 

At the base of the hill, on its southern (scarped) side, 
there are calcareous shales and thin limestones, in vertical 
position, having a strike E. 20° N. Thin beds of quartzite 
follow, divided by partings, or thin beds of purple shales, 
with gradually lowering dip, at 85°, 75°, 65°, 45° N., a few 
degrees E. -The hill itself is a mountain of almost solid 
quartzite, which, near the summit, has a dip N. 10° H. at 23°. 
The stone is softish to hard, siliceous, and, in colour, white 
to reddish. About half-way up, the quartzite contains a 
number of siliceous concretions, in the form of balls, ranging 
in size from that of marbles up to cricket balls. These have 
a rounded or flattened shape, sometimes possessing an 
equatorial ring, and are harder than the matrix in which 
they occur. The great hill is almost bare of vegetation (see 
Howchin’s ‘‘Geology of South Australia,” fig. 49, p. 66). 

A course was followed over the western shoulder of the 
hill and through a gorge on its northern side, where the 
quartzite showed a dip N. 10° W. at 27°. The path was 
followed for about 2 miles over the saddle and through the 
foot hills on its northern side. A magnificent view of the 
country lying to the north was obtained from this vantage 
ground. Immediately in front was a flat, about 24 miles 


61 


in width, drained by the Molkegna Creek, that takes its rise in 
the Patawarta Range. Beyond this river valley is a relatively 
level tableland into which the Molkegna Creek has cut, giving 
the tableland a steep scarp on its southern limits. The scarp 
shows dark-coloured beds at the top and shaly beds beneath. 
In outline, this scarp face very much resembles the “‘Reap- 
hook’’ Range (or Patterton Hill) to the southward of Blinman, 
and carries the same name from its peculiar shape. 

Still further north, at a distance of 12 miles from Pata- 
warta, the southern portions of the Angepena system of hills 
were in view. They form a remarkable circle, 8 miles in 
diameter, the dip of the rock being towards the centre of the 
area, forming a “‘pound,’ similar to the Wilpena Pound. 
Several creeks take their rise within the enclosed area, uniting 
to form the Waukawoodna Creek, which finds its outlet at 
the Waukawoodna Gap. 

The Patawarta Range appears to be greatly disturbed 
near the great hill. The dip on the southern side is vertical, 
while on the eastern side the range is broken, forming a 
jumble which passes into a bifurcation of the range; the 
northern section running east, with a few degrees south, to 
Point Well (on Point Creek), a distance of 6 miles, where 
it abruptly ends at Ann’s (“Trig.’”’) Hill. The southern 
branch trends in a south-easterly direction, and when, at about 
the same distance east as the northern branch, by a swing 
round to the southward, it converges to the nearly parallel 
Nildottie Range (or The Bunkers), so that the two ranges, 
at the point of convergence, are only separated by the Nil- 
dottie Gap through which the Artimore Creek passes. On 
the western side of Patawarta, the range curves round to 
the north-west, and then to the north-east, including Mount 
Tilley and Mount Hack, both of which are “‘trig.’’ hills, 
and continues to Angipena Head Station, a distance of 20 
miles or further. The Artimore valley widens out from the 
‘““gap” in a north-westerly direction, until due south from 
Patawarta, where it is 2 miles wide. The interval separating 
the Nildottie (Bunkers) and Patawarta Ranges is occupied 
by flags, calcareous shales, and thin limestones, with dips 
from E. to N.E. At the Artimore Head Station, situated 
within half a mile of the big Nildottie Ranges, outcrops 
show flags and purple shales with a dip N.E. at 30°. 

- For several miles, on the return track, the course was 
along the strike of the purple shales, along the Arti- 
more Creek, with the Nildottie Range on the southern 
side and the southern branch of the Patawarta Range on 
the northern. The former possesses very striking features 
—it has a dip slope of hard quartzite on its northern side, 


62 


dipping slightly E. of N. at 45°-50°. This hard back 
is underlain by softer beds, which, by weathering, cause a 
nick in the summit, and is followed by another hard quartzite 
bed, at a somewhat oblique angle to the range, which makes 
a second peak, followed by softer beds with a second nick at 
the summit. By these alternating hard and soft beds placed 
on the oblique the range is cut down, at intervals, to about 
half its height, forming a succession of house-roof structures, 
giving the range the appearance of the teeth of a gigantic 
saw. The supposed resemblance of the depressed area between 
the peaks to a succession of ‘‘bunks’’ has suggested the name 
“Bunkers.”’ Mount Lucius, a “trig.’”’ hill, is the highest 
point of the range. 

On the southern side of the Nildottie Range, the hills 
have a rounded form from the weathering of purple shales; 
they are free from trees but covered with herbage. A little 
further to the north-west, in the neighbourhood of the Wil- — 
ligon Creek (mentioned above), the quartzites form the 
southern side of the range and make a great southern escarp- 
ment. At the Nildottie Gap, where the two ranges converge, 
the rocks are much broken, with very steep dip slopes on 
the Nildottie Range, to the eastward, and a throw-up of 
calcareous beds between the converging ranges. There has 
been a contraction of the earth mass, producing folding of 
the valley beds and the bringing together of the two ranges 
at the “‘gap,’’ which has been kept open by the erosive 
action of the Artimore Creek. Strong limestones outcrop in 
the Angorigina Creek on the eastern side of Blinman. 


VII. WESTERN SIDE OF BLINMAN. 

About 1 mile from the mine, on the more northern 
road from Blinman to Parachilna, there is an outcrop of 
gneissic granite in large rounded boulders, and on the western 
side of the granite is a wide basic igneous dyke. Other basic 
intrusions are seen at intervals going west. 

About 24 miles from the mine is another outcrop of 
granite, in large spheroidal boulders, and a basic dyke, run- 
ning east and west, apparently as a continuation of the same 
line of igneous activity as that mentioned in the previous 
paragraph. The associated rocks are greatly altered. Close 
to the granite is a broken and altered dolomitic bed, which 
is intimately permeated with hematite and a little copper. 

About a mile (or little more) to the south-west of the 
above outcrops is a very large deposit of lamellar hematite 
(specular iron) in beautiful crystals mixed with siderite. The 
adjoining country rock consists of dolomitic limestones and 
flags. 


63 


As the high range, which runs in a south-west and north- 
east direction, is approached, foot hills consisting of dolomitic 
rock and quartzite flags are met with, forming the mouth of 
the gorge, and basic igneous dykes are seen on both sides 

of the road, averaging a distance of about a quarter of a mile 
apart. The one on the southern side of the road makes a 
prominent outcrop, about 30 ft. in width, but soon either 
runs out or is obscured by surface drift. The other, on the 
northern side of the road, is about 25 yards wide and strikes 
N. 20° W., and, at half a mile, crosses the road. Shortly 
before this it appears to bifurcate, the two branches with 
the sedimentary interval having a width of about 100 yards. 
Shortly before reaching the road it crosses a small creek, 
near a dolomitic limestone which has a breadth of outcrop 
of 6 yards, and is much altered by contact with the dyke. 

The gap in the ranges is about 44 miles from Blinman. 
The first definite range is composed entirely of quartzite, 
which, on weathering, breaks up into flags: dip N. 20° W. 

- at 65°. Mount Elkington is the highest point in this range. 
The next range, in the same gap, at a distance of 5 miles 
from Blinman, is more flaggy, with a dip N. 10° E. at, 55°. 


VIII. SoutH BLINMAN AND RoapD TO WIRREALPA. 


At a half-mile distance from South Blinman there is a 
low outcrop, on the right hand, consisting of flagstones with 
similar rocks forming a prominent range on the left, with a 
considerable discordance of dip in relation to each other. 
The beds on the right-hand side of the road show a dip 
S.E. at 55°, and are overlain by thick.dolomitic limestones ; 
while those on the left hand make escarpments, facing the 
west, at a dip N. 70° E. at 35°, and have a gradual slope 
to the east. 

Beyond this range, to the eastward, is a considerable 
plain, 2 miles across, with numerous outcrops of dolomitic 
rock, interspersed with shales, apparently flat, and even with 
the ground, or nearly so. The variations of outcrops on the 
plain are as follow:—Near the last-mentioned range, at its 
eastern base, are thick limestones almost even with the ground. 
At 1 mile from the range there are limestones that show a 

* kind of pseudo-vermiculate structure, with a dip of 15° E. 
At the end of the next half-mile is a ridge, 20 ft. to 30 ft. 
in height, consisting of limestones with oolitic structure, and 
includes some sandstone bands: dip E. 25° S. at 10°. For 
about another mile the road is on purple shales that form 
low exposures, the road following very nearly the line of 
strike. 

The road passes into Paddy’s Creek, at the head of which 
are flagstones and shales, with a dip of 17°. 


64 ' 


At 3 miles from South Blinman, dolomitic limestones can 
be seen on top of range, on the left hand, which pass down to 
level and cross the creek. 

At 4 miles from South Blinman, in Paddy’s Creek, a 
very thick and confused mass of dolomitic limestones and 
shales come down from the ridge to the creek, the two kinds 
of stone being crushed together: dip 55° to 90°. These 
are overlain by soft shales that are greatly contorted. Next 
follow thin quartzites separated by shale partings: dip E. 
10° N. at 45°. 

At 5 miles distance, a hill on the north side of the 
creek, about 300 ft. in height, consists of dolomitic beds, in 
part crushed. Shales occupy the opposite side of the creek : 
dip E. at 50°. 

At one-eighth of a mile further, a quartzite bar crosses 
the creek and forms escarpments on each side of the creek 
at a spot where the road finally leaves Paddy’s Creek. In 
the creek are seen indurated shales, with dip N.E. at 60°, 
while flaggy shales that show on the hill above, and overlie 
those seen in the creek, have a dip to S.E. at a low angle. This 
discordance is probably caused by faulting. 

For the next 2 miles the road follows the strike of purple 
shales, and a great development of these shales is seen in 
ranges to the southward, while to the northward the view is 
bounded by great escarpments of quartzite. 

At the 7-mile stage, the purple shales are still in evidence 
(dip N. at 55°), and the road continues on the strike of 
these beds almost to the Erengunda Creek. 

At 94 miles from South Blinman, the purple shales, on 
the southern side of the road, have a dip of 85°, and are 
faulted. 

Within a quarter of a mile of Erengunda Creek there 
is a thick dolomitic bed (not much above the general level), 
then follow purple sandstones (dip N. 65° EH. at 45°), which 
rise into a high escarpment on the eastern side of the road 
(?500 ft.). This is probably the same bed which makes bold 
escarpments seen on the northern side of the purple shales, 
as described above. 

On the northern side of the purple sandstones there is a 


_ very thick series of limestones, forming a low range on the - 


southern side of the above-mentioned creek, 90 yards wide, 
and includes a great variety of dolomitic rocks. The creek 
has cut its way through these limestones (dip 80° E. of N. 
at 55°), which cover the whole width of the creek (probably 
100 yards), and extend to 38 yards beyond, on the northern 
side, where they are overlain by calcareous shales. 

On the Wirrealpa side of the Erengunda Creek is the 


65 


‘“‘Half-way Hill’ (otherwise called the “Big Hill’). At 200 
yards up this hill there is a quarry in shales that exhibit 
‘a small syncline, or kink, in the beds, ending in a dip 
at 20°. MHalf-way up the hill the beds consist of thin 
dolomitic limestones, separated by earthy nartings, the latter 
being indurated by cementation show differential weathering, 
and stand out from the limestones in relief: dip N.E. at 40°. 
In the upper part of the hill, on the road, the dolomitic beds 
become thicker and are separated by shaly partings: dip N. 
10° E. at 42°. Near the top of the series are Archaeocya- 
thinae beds. 

Quartzites are on the northern side of the great dolomitic 
belt, and these are succeeded and intercalated with more 
dolomitic beds. 

[For descriptions of the section from the ‘‘Big Hill’’ to 
Wirrealpa, see under Section XV., as that part of the journey 
was made from Wirrealpa. | 


IX. WIRREALPA AND NEIGHBOURHOOD. 

The Wirrealpa Head Station stands on purple sandstone 
flags which underlie a series of thin-bedded limestones. The 
general strike of the country is north-easterly, with a south- 
easterly dip. Behind the station house, soft decomposing flags 
can be seen’ in the small creek, with a dip E. 20° S. at 70°, 
the beds making a curve round at the homestead. Within a 
short distance of the house, bands of oolitic limestone occur 
in calcareous and sandy shales. These bands are of much 
interest, as they include layers, up to a few inches in thick- 
ness, of broken and thickly-matted shells of Obolella, 
Pteropods, and Trilobites. These beds have every appearance 
of being shore deposits, their oolitic structure and the frag- 
mentary condition of the organic remains, closely matted 
together, all point in that direction. Associated with the 
same beds are some very fine-grained and laminated sandy 
layers, which show worm burrows and casts; the burrows are 
in the form of vertical tube-like passages, ‘while the worm- 
casts are seen on the flat surfaces of the slabs: dip 8S. 20° EH 
at 70°. Ata slightly lower position in the series is a remark- 
able layer of limestone, up to 6 in. in thickness, which is 
thickly crowded with flattened and spheroidal nodules of 
Girvanella, up to an inch and more in diameter. Their 
determination was made by thin microscope sections by which 
the typical structure of this organism was shown. Some of 
the nodules thus examined, however, failed to give a clear 
definition of structure; the very minute form of the tubes 
had, by molecular rearrangement, become more or less blended 
with the matrix. 


66 


A stronger bed of limestone, 10 ft. in thickness, under- 
lies the fossiliferous beds, just mentioned, and makes a low 
ridge that can be followed by the eye for a long distance. The 
limestone has undergone differential weathering from the pre- 
sence of siliceous material which shows in relief as blotches, 
casts, and reticulations of an arenaceous character. No organic 
structure could be detected in these objects in relief, but 
some of them strongly suggested casts of Pteropods and other 
organisms. The limestone is laminated at some levels and is 
subnodular.. Near the homestead, on its northern side, the 
bed has a dip to the S.E., from which point it gradually 
curves round to the S., then S.W., then W. 20° S. at 20°, 
then W. 20° N. at 40°, from which point it makes a strike, 
for about a mile, parallel with the eastern side of the old 
road, then makes a sharp twist in the form of the letter S. 
At 14 miles from the homestead, at a small creek where the 
road makes a sharp turn to the north, the beds dip W. 20° S. 
at 80°. About 100 yards from this point the beds are cut 
by a dip fault and the purple flags are thrown against the 
faulted face. The purple shales dip S. 20° W. at 65°. The 
limestone is thrown 53 yards to the S.W., when the dip is 
S. 20° W. at 70°. On the southern side, the oolitic lime- 
stones and Gurvanella bed that occur near the Wirrealpa 
homestead, show on the rise, with a dip W. 20° N. at 65°. 
These overlying beds are evidently faulted against the lower, 
as the strike is divergent. On the low rise situated between 
the small creek and a larger one, at this point, a great number 
of fragments of the Archaeocyathinae limestone occur, no 
doubt brought down with the alluvium of the creeks. 


X. Up a TRIBUTARY OF WIRREALPA CREEK. 


The main creek passes the station house on its south- 
eastern side, the bed of which is thickly strewn with boulders 
and gravel. Near the bottom end of small creek that is a 
tributary to the Wirrealpa Creek, on the western side of the 
“Trig’’? Hill, bleached and rotten purple shales have a dip 
of 90°. In going up the creek the dip decreases. A few 
hundreds of yards up, shelving purple sandstones, etc., are 
seen in quarry, with a dip S. at 23°. Massive purple sand- 
stones dip 8.W. at 18°. Higher up, where the creek bifur- 
cates, the right-hand branch exposes a 15-in. bluish limestone 
and a 3-ft. earthy limestone, with a shale bed between: dip 
W.N.W. at 34°. At 300 yards up the left-hand branch there 
is a bar of limestone with wavy and concentric structure 
having a dip W.N.W. About a quarter of a mile up this 
creek, several large loose stones, up to 2 ft. in diameter, con- 
taining Archaeocyathinae remains, rest on a bar of limestone, © 
but the latter does not appear to contain similar remains. 


67 


At half a mile up this creek a thick limestone with 
wavy structure occurs (dip W. 10° S.), and is overlain 
by purple shales. The latter are overlain by other 
thick wavy limestones that form a prominent hill and cross 
the creek near a group of gum trees; the dip in creek is S.W. 
At 300 yards further up another bed of wavy limestone crosses 
the creek, near the centre of which a large block of blue lime- 
stone (3ft. in length) thickly studded with Archaeocyathinae 
rested, but it had no stratigraphical relationship with the bed 
im situ. Overlying the last-named limestone are more purple 
shales, and, by folding, the same wavy limestone, underlying 
the. purple shales, is brought into the creek again, higher up. 

The creek in which the above observations were made is 
a small tributary of the main Wirrealpa Creek, which latter 
passes close by the Wirrealpa Head Station, and according to 
the pastoral map, is a continuation of the Artimore Creek, 
which takes its rise at the Patawarta Hill. Numerous loose 
examples of the Archaeocyathinae limestone were observed 
both in the wash of the small creek and on the shelving banks 
that bordered the creek. While I could not locate the parent 
rock, they are, possibly, local in their origin, as the creek 
in which they occur is only about 2 miles in length. They 
may have been derived from some of the thinnish beds of 
limestone that cross the creek, and which are much broken 
up, or, possibly, from the main Archaeocyathinae limestone 
further afield. The country, for miles around, is composed 
of purple shales, sandstones, and limestones. One piece of 
fossiliferous rock picked up in the creek was composed almost 
entirely of long lath-like organisms, the nature of which has 


not been determined. The creek will well repay further 


investigation. 


XI. Tae OBoLELLA LIMESTONE ON THE ROAD TO THE OLD 
WIRREALPA STATION. 


Followed the Blinman road for half a mile to the junction 
of the track leading to the old Wirrealpa station buildings, 
passing over purple shales, with dip W. 20° N. Following 
the old road over the first rise (low), flags outcropped, with 
a dip W. at 33°. Second low rise, thin impure limestones 
(dip W.). that are neither oolitic nor fossiliferous. Several 
such thin-bedded impure limestones occur interbedded with the 
purple shales and arenaceous flags. The strike becomes 
S. 20° W. (dip W. 20° N.). The road crosses a tributary 
of the Wirrealpa Creek in which good sections of purple 
shales and flags are seen: dip N.W. at 30°. In a low rise 
to the westward of this creek impure streaky limestones occur. 
In a second small tributary (east of the high rise) calcareous 


68 


flags strike W. 20° S. (dip S. 20° E. at 25°), and as the 
road rises above the valley a distinct fault line can be seen 
on the road that has the effect of splitting the purple shales 
which dip at a high angle: strike of fault W. 10° 8S. A 
little further on flags are seen in a washout: dip S. 20° E. 
at 35°. 

At about 24 miles from the Wirrealpa Station, a con- 
siderable rise of flaggy quartzites occurs, in which is situated 
the Wirrealpa Copper Mine. The mine, which is a small and 
new venture, is a bedded lode of shale, 2 ft. in width, lying 
between two beds of quartzite, each being about a foot in 
thickness: strike W. 10° S. (dip S. 10° E. at 40°). Over 
the rise in which the copper mine is situated the country gets 
a twist in which flags strike S. 20° E. (dip W. 20° S. at 28°). 
This rise forms a bold scarp to the west, at three-quarters 
o* a mile distant, where the beds are seen to gradually swing 
round to the strike and dip last quoted. 

On the top of the next rise on the road, 34 miles from 
the present station house, the fossiliferous (Obollela) lime- 
stone occurs. It is evidently the same bed as that which 
carries Obolella in such numbers near the head station. The 
limestone is about 5 ft. in thickness and is more solid than 
the outcrops near the house. The fossils are found mostly in 
the upper portions of the bed and are rare in the lower por- 
tions. The rock is almost completely oolitic in structure, and 
while it carries a diversity of organic remains it is particu- 
larly characterized by the presence of the brachiopod Obolella. 
Slabs can be got which are formed by one mass of the valves 
of this shell: strike of the bed W 20° S. (dip 8. 20° E. 
at 15°). . 

en 150 yards across the valley, to the northward, 
another limestone makes a prominent outcrop. This is the 
limestone which accompanies the Obolella limestone near the 
head station. It is known as the ‘‘ridge’’ limestone, or the 
‘“‘5-mile ridge,’’ as it makes a prominent feature across the 
country for 5 or 6 miles. This ridge limestone and the 
Obolella limestone cut the road at a right angle. 

Following the fossiliferous limestone westward, the beds 
cross the Wirrealpa Creek within a few hundred yards of the 
road, with a strike W. 20° S. and dip S. 20° E. at 20°. 
Shortly after crossing the great creek the beds swing round 
sharply to S. 20° W. and dip E. 20° S., passing under the 
escarpment of red sandstone, mentioned above, which is the 
same range in which the copper mine is situated. 

Returning to the road I followed the outcrops of the 
fossiliferous beds in an easterly direction. Here the dip 
swings round to the south, then S. 20° W., then S.W. at 23°. 


69 


At about half a mile from the road the ridge is suddenly 
broken and twisted over a distance of 150 yards, when the 
beds again form a ridge as an isolated hill: strike N. 20° W. 
(dip W. 20° S. at 28°). The ridge is maintained for about 
200 yards, when it ends abruptly—cut by a fault—while the 
limestones are thrown at nearly right angles to their former 
direction: strike E. 10° S. (dip at 35°). So far as the beds 
could be followed, by sight, they continue on the same strike ; 
then, at a mile distant, they appear to curve round towards 
the east, or some point south of east. _ 

Starting again from the Obolella bed that crosses the 
road to the old station, 34 miles from the present Wirrealpa 
Station, going north, a limestone ridge forms the foot hills 
of the main escarpment range, and has its strike in the 
direction of the old station, and appears to be faulted. A 
very much brecciated limestone occurs between this ridge 
and the great Archaeocyathinae limestone which outcrops in 
the creek, a little higher up, where it is exposed in great 
spheroidal masses. The brecciated limestone {which shows 
bedding planes) has a dip W.S.W. at 50°, and contains 
angular fragments of purple quartzites, etc. 


XII. THe Otp WIRREALPA STATION. 


At the old Wirrealpa station (7 miles north of the present 
_ station) there is a great show of the Archaeocythinae lime- 
stone series, making 100 yards of outcrop: dip W.S.W. at 55°. 
There is an extraordinary break up of the rocks which are 
seen in the Wirrealpa Pass. The great quartzite range, which 
forms a high peak at Wirrealpa Hill, comes to an abrupt 
end near the old station buildings, and the beds in the valley, 
as well as much of the thick limestone on the opposite side, 
have been reduced to a breccia. Towards the centre of the 
valley the beds are mostly quartzitic, and are so crushed that 
they form a fault rock of great width. The great quartzite 
hill disappears—cut off by a fault—on the north-eastern side - 
of the station ; the limestone on the opposite side of the valley 
forms a conspicuous peak, which can be seen from a great 
distance. 

At the old station the beds dip to the 8.W. Following 
_ up the creek the beds make a curve. At half a mile distance 
they dip W. 10° S. at 35°; then a little further, the lime- 
stone gives a reading of W. 20° S. at 75°, which has the 
appearance of being almost at right angles to the dip of a 
conspicuous hill (marked ‘‘First Hill’? on map), with lime- 
stone near its summit, situated in a direction to the 8.S.W. 
As near as could be determined, the beds of this distant hill 
have a dip W. 20° N. at 40°. 


70 


The country gives every evidence of great and conflicting 
earth movements. The Wirrealpa Hill, which forms the 
terminal point of the Mount. Lyall Range (which extends in 
a north-easterly direction), is the exact counterpart of the 
Parachilna quartzite escarpment. The dip of this hill is 
also on the curve. The southern side of its termination dips 
south-west ; whilst, on the opposite side, the dip is W. 20° N. 
The Archaeocyathinae limestone overlies the quartzite at the 
base of the hill (as it also does in the Parachilna Gorge): 
dip in the centre of the curve is W. at 45°.. The last ex- 
posure of the quartzite, in the big creek, on its eastern side, 
has a dip S.W. at 40°. 

The Archaeocyathinae beds have some features of special 
interest. In one part the rock is of a light colour and very 
pure, and although the larger forms of organic remains are 
somewhat scarce at this horizon, those that are there are 
well preserved, and the rock mass is largely made up of small 
sponge spicules, which can only be detected in microscope pre- 
parations and do not admit of further determination. 


XIII. Visit to Mount CHAMBERS CREEK. 


Mount Chambers is an important “trig.’’ hill situated 
about 23 miles to the south-eastward of Wirrealpa. The inter- 
vening country is mostly low. Mount Chambers Creek comes 
in from the north-west and penetrates the mount, where it 
makes a bold and rugged: gorge with nearly vertical faces. 
The mount has a limestone cap, formed into a shallow 
syncline. The lmestone is somewhat impure, resting on 
purple shales, which show a sudden increase of dip. A fault 
is probably present, as the rocks, extending over a large out- 
crop, are in the condition of breccia. In one of the outcrops 
the shales have a bluish colour: dip W.N.W. at 27°. The 
limestone underlies thick purple quartzites, which can be 
seen in the gorge, and also in a small tributary creek that 
flows into Chambers Creek. In this small tributary there 
is a remarkable amphitheatre in the rocks, almost hid from 
view, in which is a spring of good water, and the walls, 
above the height where a man could reach, are well covered 
with native drawings, cut into the rock faces: dip W.N.W. 
at 24°. At a lower stage in Chambers Creek, about 2 miles 
lower down than Mount Chambers, is an important outcrop 
of the Archaeocyathinae limestone. 


XIV. Mount Lyatt. 
Travelling from the present Wirrealpa Head Station, 
by the Tea-tree Well road, at half a mile distant beyond 
the gate which admits to the Woolly Paddock, a small lime- 


71 


stone is exposed; it has a laminated structure and is under- 
going change to manganic and ferric oxides: dip S. 10° W. 
at 30°. 7 3 : 

The Mount Lyall Range extends from the old station, 
in a north-easterly direction, for a distance of about 3 miles. 
At Mount Lyall (fig. 2) it forms a sharp angie with a range 
that comes down from the north. At the angle is a prominent 
hill, about 150 ft. in height, which shows a scarp face to the 
south-east. This hill is a solid mass of limestone, brownish 
in colour, dolomitic, and very compact. The section repre- 
sents the lower beds of the Archaeocyathinae series. These 
beds follow the strike of the northern range of hills, but they 
appear to run out to the northward. It is probable that 
the scarp face of the hill is a fault plane. The hill just 
mentioned has a dip at the summit N. 10° E. at 15°. From 
the top of the hill it could be seen that the beds on the north 


4 Cy AM 
ett as ° 
I] <= 
Cig a aie Caan pe 
ee ee oS Me Lyart 
w ta] 4) 
see AES 
S > RG Ry aa UT, 
SOO eon : cy - 
CA REE : 
XX S WS 
x SS 
QS: ~ 


Y 
Y 

iy 

Ut 
SY 


Fig. 2. Geological Section of Mount Lyall. 


side of Mount Lyall were much disturbed, dipping at various 
angles both to the Mount Lyall Range and also to one another. 

The peak of Mount Lyall dips N.W. at 60°. The north- 
eastern end of the range is broken by (?) two faults. Mount 
Lyall peak dips as above, then on the eastern side the dip 
somewhat suddenly changes to nearly horizontal. After a few 
hundred yards at this level there is a sudden break, when the 
purple shales, much crumpled, are thrown down to the face 
of the quartzites with a dip N. 20° W. at 35°. A crush-zone 
occupies the fault area for a thickness of about 9 yards.. A 
series of underlying dolomitic limestones, purple shales, and 
quartzites follow with a rising dip, going east, to 50° (see 
fig. 2). Just round the northern end of the range there is 
an oblong dyke of intrusive rock. On the eastern side of 
the dyke there is a crush-rock, composed of flaggy shales, of 
considerable extent. This is probably on the line of fault, 
noted above, that cuts through the north-eastern part of the 
range. 

There are low exposures of the Archaeocyathinae lime- 
stone, in the form of a fragment (probably hmited by two 


72 


fault planes), with a strike W. 20° S., and this is cut off by 
a fault that throws the quartzites against the limestone in 
an oblique line. Nearer the foot of the range the lower beds 
of limestone in the series occur, having a dip N. 20° E. at 
about 25°. 

A little south of this, cut off by faulting, is a small 
quarry in which the decomposed sandstone is seen, on one 
side, having a dip W. 10° N. at 15°, and is thrown down, 
on the eastern side of the quarry, with a dip E. 20° N. 
at 75°. The strike of the fault, which cut off the Archaeocya- 
thinae limestone, seems to have a bearing W. 20° N. The 
end of the limestone, where faulted, shows metasomatic change 
to iron and manganese oxides. On the western side of the 
fault are purple shales (greatly contorted), thin limestones, 
and flags. 

The general section of the Mount Lyall Range closely 
resembles that seen in the Parachilna Gorge, but is more 
disturbed than the latter. 


XV. From WIRREALPA TO THE ‘‘Bic Hi1uu’’ 4) on THE Roap 
TO BLINMAN. 


At 4 miles out from the station (after having passed 
over a flat of purple shales and flags, with an occasional thin 
bed of impure limestone), the red sandstone of the scarp hill, 
near the Wirrealpa Creek, comes down to the flat and crosses 
the road: dip S. at 15°. The road runs on the strike of these 
beds for a mile, then the sandstones swing round to south-east, 
with an increase in the angle of dip, and are underlain by a 
series of limestone bands that have an oolitic structure, which 
measure, in the creek and sides, about 50 yards of outcrop: 
dip E. 20° S. at 80°. These oolitic limestones are the same 
as have been spoken of above as the Obolella limestones. 
Within a few yards they are followed by the ‘ridge’ lime- 
stone, which forms a kind of rampart 12 ft. high: dip E. 
20° S. at 65°. On the northern side of these beds, there 
are soft decomposed flags and shales, which are perpendicular, 
or, for a few yards, reversed, to W. 20° N., with a decreasing 
dip, and then back again to easterly. A creek runs in the 
form of a loop between these outcrops. 

At 6 miles out from the station there are two conical 
hills, situated on the western side of the road, which are 
either igneous dykes, chimneys, or (?) sheets. The highest 
is probably 300 ft. in height, and con. .ts of a dark-coloured 
basic rock; but whilst it passes to the top of the adjoining 


(4) This is quite distinct from the ‘‘Big Hill” which occurs 
between Parachilna and Blinman although known, locally, by the 
same name. 


ee 


73 


hill, separated by a deep and narrow valley, no continuity 
can be traced between the two intrusions. Indeed, stratified 
beds of limestone and decomposed shales can be traced almost 
uninterruptedly across the dividing area. The beds are much 
disturbed in their strike and angles of dip adjacent to the 
igneous rock. There is a curious, and not easily explained, 
relation between the igneous rock and the sedimentary 
deposits. A limestone appears to go up to the centre of the 
igneous dyke (or (?) sheet), in the centre of the line of strike. 
The limestone has a dip EH. at 70°, whilst the strike of the 
igneous rock is N.E. 

Separated by a belt of rotten shales and quartzite flags, 
100 yards in width, is another igneous dyke (or (?) sheet), 
near to the road. Its composition is very distinct from those 
previously mentioned, although on about the same line of 
strike. At the north-east side of the hill there is a bold 
outcrop of rock consisting of limestone and shale, closely 
adjacent to the igneous rock. 

Following these igneous outcrops, in a westerly direction, 
is Sandalwood Flat, about 1 mile in length, in which the 
rocks continue mostly on the same tine of strike. At the 
northern end of Sandalwood Flat, and on the same side of 
the road as the preceding, another igneous rock is seen in a 
steep cliff in a creek. It has the same strike (N.E.) as the 
smaller igneous hill, next the road, described above, and 
possesses a similar rock texture. 

A second ridge, on the westward, rises to about 300 ft. 
in height. The lower half of the hill consists of a hard 
laminated and dark-lined quartzite: strike E. 20° S. at 90°. 
The upper half of the ridge is a very close-grained igneous 
rock, with features distinct from the two other varieties 
already mentioned. 

Near the base of the ‘‘Big Hill,’’ on its eastern side, 
an important basic igneous dyke is exposed on the road. Its 
strike is, apparently, north and south, and is 34 yards wide. 
It occurs in dolomitic limestone, which latter is somewhat - 
altered by contact with the intrusive dyke, and contains some 
copper ores, as well as a very large mass of ferrugineous and 
copper- -stained quartz. 

There are thus five important igneous intrusions near 
together and adjacent to the road on the Wirrealpa side of 
the ‘Big Hill.”’ I very much regretted that no opportunity 
presented itself for -’ aking a more detailed examination of 
this very interesting ~ igneous field. 

As the “‘Big Hill” is approached from the eastern side, 
thick limestones appear on the eastern side of the road: dip 
E. 20° N. at 15°. A sandstone occurs in the creek, with a 
dip W. 20° S. at 55°. It is, apparently, included in the 


74 


thick limestones, or, possibly, is nipped in by a fault; the 
beds seem to bifurcate about the spot. The limestone ex- 
posed on the eastern side of the road has the appearance of 
a shallow syncline. The great mass of limestone that forms 
the main part of the ‘‘Big Hill’? comes up from the Eren- 
gunda Creek (see under Section VIII.). In its purest por- 
tions it forms a white and grey marble (dip N. 30° E. 
at 30°), with remains of Archaeocyathinae. The best fossili- 
ferous horizon is in the upper beds, near the public road. 

[The observations at this point join on to those given in 
the traverse from Blinman to the ‘‘Big Hill,’ in Section 
VEE] 


XVI. Visit To THE GRINDSTONE RANGE, BALCORACANA 
CREEK, AND THE WILKAWILLINA GORGE. 


This trip was in a southerly and south-westerly direction 
from Wirrealpa Station. Going in a south-westerly direction, 
the track was over flaggy sandstones. At 1 mile distant, 
in small creek, soft sandy flags were exposed, showing false 
bedding: dip W. at 25°. At 3 miles out, descending to a 
valley (three-quarters of a mile in width), the exposures 
were still sandy flags, much false bedded: dip E. 20° S. 
at 35°. In a dry .creek, near the centre of the valley, 
sandstone and flags have a dip EH. 20° N. at 35°. After 
crossing the valley and ascending a small rise, I passed over 
into the Balcoracana Creek. This creek, which is the most 
important waterway in the neighbourhood, takes its rise in 
The Bunkers, which are a continuation of the Nildottie 
Ranges that occur on the southern side of Patawarta. There 
was a strong flow of water in the creek at the time of passing. 

On the southern side of the Balcoracana Creek is situated 
the Grindstone Range, or the ‘“‘Little Bunkers.’’ These form 
an isolated range of hills, about 3 miles from Wirrealpa, and 
are intersected, at one end, by the Balcoracana Creek. They 
consist of purple sandstones and shales, the latter, wasting 
more rapidly than the former, produce an outline of peaks 
and depressions, from which feature they have received the 
name of ‘‘bunkers,’’ on account of their resemblance to the 
main range of The Bunkers, on their western side. The 
‘“‘white cliff,” or the ‘‘grindstone cliff,’’. in the range, is 
formed of a sharp fine-grained sandstone, which is used locally 
for making grindstones, from which the range has received 
its secondary name: strike S. 20° W., dip easterly at 63°. 

Went up stream, in the Balcoracana Creek, to the junction 
of small creek which comes in from the north-east. The 
main creek channel, almost immediately, going up stream 
turns due west and makes a gorge that penetrates the high 


75 


range (The Bunkers), which has a south-south-easterly direc- 
tion from the “Big Hill’ of Erengunda Creek to the locality 
under examination. 

Having reached the base of the high range, the eastern 
side of the range was found to form a dip slope of the 
Archaeocyathinae limestones, from top to bottom, but the 
fossils are not very well preserved. The dip of the beds, at 
the bottom of the range, reads H. at 48°. The limestone 
beds have a rolling curviture along the strike which, at times, 
causes angles in the outcrops, with slightly varying directions 
of dip. I was informed by Mr. Napier that the range has 
a steep slope on its western side and is followed by a high 
range of hard quartzite rock, similar to that which accom- 
panies the Archaeocyathinae beds at Wirrealpa Hill, at the 
“Big Hill,’’ on the Erengunda Creek, and elsewhere. These 
quartzites, with their interbedded shales, form the true 
‘“‘Bunkers.”’ 

In a small creek, on the eastern side of the range, there 
is a thin bed of laminated limestone that is extremely con- 
torted, making acute v-shaped folds, with a dip S.W. at 45°. 
This bed is similar to the perpendicular and contorted thin 
limestone met with, in about the same stratigraphical posi- 
tion, at the old Wirrealpa station. A quartzite overlies the 
limestone (as it does at the old station), then follow rotten 
purple slates, extending over a distance of half a mile, between 
the base of the range and the Balcoracana Creek. 

At the southern side of the Balcoracana Creek, facing 
the main range, is a high hill showing a scarp face to the 
valley. The beds, seen in section, consist of thick, soft, and 
red-coloured sandstones, interbedded with purple and other 
coloured, thin-bedded, argillaceous beds, which cross the creek 
near the east and west bend of the stream. 

Resting on the dip-slope of the last-named red sandstones 
and shales is the bed corresponding to the Five-mile Ridge 
limestone, which, at a distance of a few yards, is followed 
by the oolitic and Obdolella limestone (see Section). At the 
immediate junction of the north-easterly tributary (mentioned 
above) with the main creek, a fine and complete section of 
these limestones occurs in the creeks, with a dip E. 10° N.. 
at 45°. The fossiliferous limestone is sometimes flaggy and 
nodular in structure. ‘The bed is rich in Obolella and other 
Brachiopods (in some cases the interior of the shells is filled 
with rhombohedral crystals of calcite), Pteropods, and 
abundant fragments of Trilobites, but none were seen suffi- 
ciently complete to permit of further determination. The 
fossiliferous zone, so far as a few minutes’ examination could 
determine, was limited to a few inches in thickness. 


76 


Overlying the above-mentioned limestones is a_ thick 
series of soft, thick-bedded, red sandstones, interbedded with 
sandy shales, usually red coloured, with dip E. 10° S. at 48°. 
These beds are intersected by small creeks, which have carved 
out fair-sized hills on the western side of the main creek. 

In superior position to the last-named red sandstone is a 
limestone, 5 ft. wide, laminated and contorted and inter- 
bedded with purple shales, having a dip E. 20° N. at 65°. 
Then follow, in ascending order, purple and greenish shales 
with thin bands of limestone, then a series of small ridges 
showing scarp faces to the north-west, consisting of red sand- 
stones and flags. A high ridge follows before reaching a 
valley which separates the latter from the Grindstone Range, 
or Little Bunkers, as described above. 

The north-eastern angle of the Grindstone Range was 
then followed, where the range passes down to soft and decom- 
posing sandy flags and shales, which cross the Balcoracana 
Creek on that side: dip E. 20° N. at 50°, changing to a 
dip EH. 

WILKAWILLINA GORGE. 
This gorge occurs in the Mount Billy Creek (or Ten- 


“mile Creek), situated about 6 miles to the southward of the 


Grindstone Range. A remarkable exposure of the Archaeo- 
cyathinae limestone occurs at this spot. The limestone is 
associated with a great range of hills that are about 600 ft. 
in height: strike N.W. (dip S.E. at 15°). The fossiliferous 
limestone forms the bed of the creek for about a quarter of 
a mile in length. Near its upper part the rock is almost 
one mass of Archaeocyathinae. The matrix, as a whole, is 
a white crypto-crystalline marble which, throughout the 
greater thickness of the limestone, only occasionally shows 
the presence of the fossils; the latter, most likely, having 
been largely destroyed in the alteration of the rock texture, 
but near the top the fossils are better preserved. The most 
striking feature of this outcrop is that the grain of the stone 
permits its ready fracture, in such a way that the fossil ‘‘cups’’ 
can be broken out from the matrix so as to show the external 
form of the organism. This is the only instance that has 
come under my observation in which this can be done. The 
matrix in which the Archaeocyathinae are usually included 
is of an amorphous and refractory character, and is of the 
same nature within the fossils as in the surrounding matrix, 
so that the rock fractures uninfluenced by the presence of the 
organic remains. The only approximate condition for obtaining 
the objects free from the matrix, naturally, in the normal 
limestone, is where the organism has undergone silicification, 
by which the fossil is produced in relief on the weathered 
surface, as in the case of the Ajax specimens. 


17 


The fossiliferous limestone in the Wilkawillina Gorge is 
overlain by purple shales, and underlain by strong beds of 
purple sandstone, divided by thinner beds or partings of the 
same kind. 


XVII. LirHoLtocic FEATURES. 


The country dealt with in this paper supplies the most 
extensive series of the Upper Cambrian beds that has come 
under my observation. The lithology of the beds agrees very 
closely with the occurrence of beds of the same age in other 
parts of South Australia, and include quartzites, sandstones, 
shales (or slates), limestones, and intrusive igneous rocks. 

The Quartztes are of two kinds. (a) A light-coloured, 
very fine-grained, and siliceous rock that possesses great 
resistance to weathering, and forms pointed and serrated 
outcrops that- make prominent features in the landscape. A 
rock of this kind usually underlies the Archaeocyathinae lime- 
stones. (6) The other variety is of a dull-red or purple 
colour, and is usually divided up into layers of a few inches, 
or a foot or two, in thickness, separated from each other by 
indurated shales or finely-laminated bands of quartzite. The 
term ‘‘flaggy’’ has been used in the present paper to describe 
features of this kind. 

Sandstones occur of various colours, mostly red. These 
are especially characteristic of the eastern side of the ranges. 
They are, usually, more or less argillaceous in composition, 
finely-laminated, and cross-bedded, generally soft, and some- 
_ times friable. They have been utilized, to some extent, as 
flags; but are, generally, too soft for such a purpose. 

Shales (or (?)Slates).—These form the predominant 
element in the sedimentary rocks of the district. In some 
instances they may have developed an incipient cleavage and 
could be called slates; but, in a general way, they are readily 
fissile, splitting on the bedding planes, and from intimate 
jointing break up into cuboidal fragments. As it is not always 
an easy matter to draw a line of distinction between slates 
and shales in the field, the term ‘‘shale’’ has been adopted, 
uniformly, for this class of rock throughout the paper. They 
sometimes possess a greenish or drab colour, but they are pre- 
ponderantly of a purplish tint and are, collectively, classed 
as “purple shales.’”’ Like the quartzites and sandstones, they 
- are often divided up into definite layers of a few inches 
thick and have the features of ‘‘flags.’’ Occasionally they 
make prominent heights, but more commonly they weather 
rapidly and make low ground. 

The Limestones are both numerous and of varied types. 
Magnesia enters largely into the composition of many of 
them. Some are true dolomites, while many others have a 


78 


marked dolomitic character. As it is often impossible, in 
the field, to distinguish a true dolomite from some dolomitic 
limestones, it was considered better to describe this class of 
rock, as a whole, as dolomitic limestones. There are bluish 
limestones, exactly similar in appearance, to the Carboniferous - 
limestones of Europe; siliceous limestones, and arenaceous 
limestones. Many of the thinner limestones have an oolitic 
structure, and it sometimes happens that the oolitic grains 
and rounded sand grains occur together in a rock in about 
equal quantities. In one instance (in Wilkawillina Gap) an 
oolitic limestone, at one particular zone, had become altered 
to an oolitic flint. The Archaeocyathinae limestones form a 
group by themselves. They are, usually, relatively pure, but 
in places show siliceous and earthy veins and patches, which 
weather into relief. The same happens when the included 
fossils have undergone some measure of silicification, when they 
make most interesting and showy faces on the weathered sur- 
face of the rock. Occasionally the limestone partakes of the 
nature of a marble, either dark coloured or nearly white. A 
change of texture, of this kind, is generally destructive of the 
organic remains, which become altered and indistinguish- 
able from the cryptocrystalline matrix. The Archaeocya- 
thinae outcrops in the district appear to be limited to two 
distant localities, the one on the western side of Blinman and 
the other on the eastern. In the western outcrops the beds 
form the foot hills of the Flinders Ranges, facing the western 
plains, where they extend both north and south of the Para- | 
chilna Gorge. In the eastern areas of outcrop they are more 
irregularly placed. They follow, to some extent, The Bunkers, 
running in a south-eastern direction, where outcrops were 
visited at the “‘Big Hill,’ at the head of the Balcoracana 
Creek, and in the Wilkawillina Gorge, measuring from point 
to point a distance of 16 miles.. There is, apparently, another 
line of strike that passes in a north-easterly direction, 
diverging from the ‘“‘Big Hill,’’ passing by the old Wirrealpa 
station; a small faulted patch occurs near Mount Lyall; 
and then, following in the same direction (after a distance of 
about 20 miles), there is another important outcrop of the 
limestone in the Mount Chambers Creek. 

The Igneous Rocks are restricted, so far as the present 
observations are concerned, to two localities; one of these is 
in the neighbourhood of Blinman (especially developed on the 
western side of the township), and the other occurs on the 
eastern side of the “‘Big Hill,’’ nearer to Wirrealpa. The 
occurrence of intrusive pipes, of a circular outline, are inter- 
esting features as indicating ancient volcanic vents that have 
been cut back by denudation. Petrographically, the rocks 


79 


belong, mostly, to the diabase types (see Benson, ‘‘Basic 
Rocks of Blinman,’’ Roy. Soc. 8. Austr., xxxiii., 1909). The 
amount of alteration induced in the adjacent rocks by con- 
tact metamorphism varies considerably with the different 
dykes. In some cases, little or no alteration can be detected, 
while in others some quartz and allied minerals appear to 
owe their development to contact with the dyke. Siderite, 
hematite, limonite, and copper ores are frequently found in 
dolomitic limestones near the junction of these beds with an 
igneous dyke. It is an interesting fact that on the south- 
eastern spurs of Mount Remarkable a small field of igneous 
dykes occur, having similar petrographical features to those 
found near Blinman, and which intrude rocks of a similar 
geologic age (see Thiele, Roy. Soc. S. Austr., xl., 1916, 
p. 580). 

The association of igneous dykes, dolomitic limestones, 
and copper ores is a characteristic feature of the Blinman 
mining field. Dr. Lander’s mine, which I visited, situated 
near to Blinman, about a quarter of a mile to the southward 
‘of the old Blinman to Parachilna road, is typical of most 
of these mines. The ore occurs in bunches, immediately 
below an igneous dyke, which is intrusive at a low angle. 
The carbonates and oxides of copper, siderite, and hematite, 
together with some quartz, are distributed through a metalli- 
ferous zone, mixed with vein stuff, up to 9 ft. in diameter. 
Below the ore zone is a'broken-down shale, containing a little 
ore, and this latter rests on dolomitic limestone. 

The Scenic Aspects of the Flinders Ranges are often 
very striking, and certainly unique in South Australian 
scenery. Under arid conditions, the weathering agencies have 
sculptured the country into sharp and rugged outlines, pro- 
ducing bare hills and mural cliffs. The prevailing red colour 
of the rocks also gives an unusual tone to the landscapes and 
lends itself to uncommon colour effects. When standing on 
some vantage point, commanding a view of the surrounding 
hills, especially with the slanting rays of the setting sun 
thrown upon the scene, the picture is full of a weird beauty. 
The bare scarps of the ranges look like enclosing walls, the 
red rocks possess a higher colour by reflecting the lurid sun- 
set, and the glow on the rising mists of the valleys all combine 
to ‘give the appearance of a vast furnace or the floor of a 
smoking volcano. 


XVIII. Tectonic PHENOMENA. 
The main geological structures of the region under 
description are relatively simple. The great fault-scarp, 
facing west, by which the Flinders Ranges are suddenly cut 


80 


off in that direction and are replaced by the flat 
and sandy shores of Lake Torrens, makes a very 
sharp boundary line, both topographically and 
geologically. 

At the entrance to the gorge the Archaeo- 
cyathinae limestones are at a lower level and 
considerably lower angle of dip than the great 
scarp quartzite on which they rest. This strati- 
graphical discordance may possibly represent 
an angular unconformity in the geological suc- 
cession, separating two series of beds that are 
of different ages. Its evidence in this direction 
is rendered doubtful, however, in that the 
displacement (if such exists) occurs on the 
nearly vertical face of the escarpment which 
forms the eastern wall of the great South Aus- 
tralian rift valley. This line of major faulting 
is accompanied by many secondary fractures and 
block faulting, the respective ‘‘blocks’’ settling 
down at various angles. It was a matter of 
regret that the necessary time could not be 
spared for making such detailed observations as 
might have settled this interesting question. 

The tectonics of the ranges are fundament- 
ally based on periclinic and cycloclinice fold- 
ings, together with much lateral movement. The 
long curves of the folds canbe noted from the 
train, going north, from Mernmerna Station 
(fig. 3). At the latter position, the western 
scarps of the Elder Range make a bold feature; 
and a little further north, the western wall of 
the Wilpena Pound Range is equally impressive. 
A rough sketch of these mountains in section is 
given here. 

Blinman occupies the centre B a great 
earth movement of elevation. The tangential 
forces have acted from all sides, almost equally, 
with the effect that the whole district, from 
Parachilna Gorge to Wirrealpa Old Sta- 
tion, in a diameter of 20 miles, has been 
raised in the form of a vast dome. The upper 

b 
. 
| 


Mernmerna 


WilpenaPound 


Fig. 3. 
A Diagrammatic Sketch-section from Mernmerna to Blinman, 40 miles. 


beds form the outer rims of the dome, and the 
centre, much reduced by denudation, exposes 
the lower members of the series. According to 
official figures, Blinman has a height of 
2,020 ft. above sea level. Parachilna railway 
station, on the plain, is 465 ft. above the sea 


Wirrealpa 


81 


level, and the great escarpment at the Parachilna Gorge is 
probably 1,000 ft. higher than the railway. 

The quaquaversal dip of the beds around the apical 
portion of the dome is fairly consistent as to direction, but 
varies much in the angle of dip. The Archaeocyathinae lime- 
stones represent the highest exposed horizon on the western 
side, the beds in superior position having been thrown down 
by the great north and south fault, and have become obscured 
by recent alluvia and blown sand. From the gorge on the 
“west to Blinman, a distance of about 10 miles, the prevailing 
dip is westerly. From Blinman to the Archaeocyathinae beds 
at the old Wirrealpa station, on the east, at a similar distance 
of about 10 miles, the prevailing dip is easterly. At about 
the same distance to the northward of Blinman is Patawarta 
Hill, which probably represents the thick quartzites that 
underlie the Archaeocyathinae limestones at the Parachilna 
Gorge, and, if so, those fossiliferous limestones might be 
expected to occur on its northern slopes. 

A 10-mile section of rocks that lie in one direction, at a 
fairly high angle of dip, would make a very thick series, and 
suggests the possibility of faulting along the strike that might 
cause a repetition of the beds and give a fictitious appearance 
as to their thickness. Minor faults were recognized in several 
places, but nothing came under my observation that looked 
like a repetition of the beds on a large scale, although in a 
single traverse such an occurrence might easily be overlooked. 

One of the features of the district is the great amount of 
crush-rock that is developed, at intervals, over scores of square 
miles. This class of rock takes all the forms usually developed 
under such conditions, v2z., crush-breccia, crush-conglomerate, 
over-riding, and sometimes telescoping, when the shales and 
dolomites interpenetrate one another. Some of the purple 
shales are, in places, crushed into a confused mass in which 
signs of bedding can only be recognized in disconnected frag- 
ments. These features are, perhaps, in greatest evidence 
where the igneous rocks are in close proximity. The only 
locality where I have noticed autoclastic phenomena in any 
degree comparable to this is along the flanks of the great horst 
that forms Mount Remarkable (see Howchin, ‘‘Geology of 
Mount Remarkable,’ Roy. Soc. S. Austr., xl., 1916, p. 545). 
In the latter case the crush-rock has been caused by vertical 
faulting on a large scale; in the northern Flinders Ranges, 
lateral faulting, by a sliding horizontal motion, has been of 
common occurrence, and would, probably, be more potential 
in causing “crush’’ than vertical movements. 

The beds above the horizon of the Archaeocyathinae lime- 
stone, which outcrop near the Wirrealpa Station and in the 
upper part of the Balcoracana Creek, are the highest members 


EE EE EE Ee —eeee OEE ES EEE ESE oe 


. 


82 


of the Upper Cambrian Division that have been hitherto 

recorded in South Australia. With the exception of a thin 

bed of laminated and contorted limestone that occurs a little 

higher in the series than the Archaeocyathinae horizon, these 

top beds are not much disturbed. They consist, mostly, of 
softish and highly-coloured sandstones and shales with one 

highly fossiliferous horizon (the Obolella limestone), which is 

of no great thickness. 

As to the physical conditions under which the beds were 
laid down, the evidence seems to point to shallow water, if not- 
dry land conditions, at some horizons. Some of the limestones 
have a nodular, or subglobular, kind of structure, which 1s 
seen on the weathered surface, and when split by the hammer 
break up into more or less rounded fragments, which have a 
close likeness to the surface concretionary travertines that form 
in calcareous soils under an arid climate. A specimen picked up 
near the old Wirrealpa station showed what had been, in the 
first instance, projecting cups of Archaeocyathinae, and then 
were contemporaneously surrounded by concretionary limestone; 
such as might have been formed following on the elevation of 
a reef of these organisms above the level of the sea. The 
very common occurrence of oolitic limestones, and oolitic sand- 
stones in which the oolitic grains and rounded sand grains are 
mixed up together (very much as they occur in present-day 
deposits laid down in a shallow lake, near Robe, which is 
alternately wet and dry) (see Howchin’s ‘‘Geology of South 
Australia,” p. 176, figs. 152-154). Still further, the red and 
friable sandstones, much cross-bedded, near the top of the 
series have features that favour the idea of a terrestrial origin. 


DESCRIPTION OF PLATE IV. 
Fig. 1. Geological sketch-section of outcrops from the mouth 
of the Parachilna Gorge to the vicinity of Bliinman. 
Fig. 2. Geological sketch-section from Blinman to the 
northern side of the Erengunda Creek. 
Fig. 3. Geological sketch-section from The Bunkers to the 
Eastern Plains. 


Note.—The Geological Sections, as described above, are made 
as detailed as the nature of the outcrops permitted. The rapid 


. changes, in succession, of quartzites, shales, and limestones, within 


short distances, render it impossible to note such occurrences, in 
detail, within the limits of the scale adopted. The dip of the 
beds, also, varies greatly, in places, within short distances. It 
must therefore be taken for granted that the sections are, to a 
large extent, generalized rather than exact. A further difficulty 
arose from the quaquaversal curves in the dip, so that in some © 
parts of the section the line follows a true direction of dip, while, 
in others, it approximates to the line of strike, in which case 
important beds, situated on one, or other, of the sides of the 
section, and running parallel with it, could not be shown in section. 


83 


Two NEw SPECIES OF LYCOSA FROM SOUTH AUSTRALIA. 


By R. H. Puiieine, M.B., C.M. 


[Read November 10, 1921.] 
PuaTE V. 


Up to the present seventy species of Lycosa have been 
described as Australian. This is probably only a fraction ot - 
the whole of this immense genus existing on our continent. 
Owing to the great powers of locomotion of the young Lycosa 
it is not safe to view every Australian species as endemic 
without further investigation. Some eremeian species also 
show variations in colour of almost specific value; but con- 
necting variations can be found which even invalidate their 
varietal value. The two species described are certainly new, 
and the types are preserved in formalin in the collection of 
the South Australian Museum. 

I have found that the species of Lycosa described by 
H. R. Hogg (P.Z.S. Lond., 1905, vol. ii., p. 569), and pre- 
served in the South Australian Museum collection, are, from 
long immersion in alcohol, in poor condition for identification. 


LYCOSA SKEETI, 0. sp. 


Q. Cephalo-thorax light brown, clothed with silvery- 
grey hair; a darker brown median streak with four similar 
streaks on each side. 

Mandibles concolorous, clothed with long silvery hair. 
Lip maxillae and sternum dark brown. 

Abdomen light brown above, dark brownish-black below, 
spinnerets of the same colour, lighter in shade. 

Legs and palpi the same colour as the thorax. They are 
clothed with fine silvery hairs interspersed with strong black 
spines. 

The eye area is prominent, and the arrangement of the 
eyes, which are black and shining, is of the ordinary Lycosa 
type. In the eye area and on the clypeus are strong, erect, 
yellowish-brown hairs. 

The markings of the dorsum of the abdomen are as 
follow :—Posteriorly, two nearly straight black parallel lines 
meeting at their ends; anteriorly to this, three parallel sinuate 
lines; in front, two lateral black, forked lines, not meeting 
medially. 


84 


Epigyne small, shining brown of simple form, v2z., two 
depressions with a median ridge. 

Total length, 65 mm.; thorax abd., 25 mm. 

This striking species of Lycosa was sent from Wilson, 
Flinders Range. 

Type in South Australian Museum. Male unknown. 

Named after Mr. H. C. Skeet, of Melbourne, an 
enthusiastic collector for other naturalists. 


Lycosa PERINFLATA, 0. sp. 
Cephalo-thorax broad, compressed, nearly circular in 


. outline; warm reddish-brown, covered with fine white 


adpressed hairs. 

Median brown lines extending on to eye area in front, 
uniting behind and then spreading into a broad fork with 
radiating brown lines and spots on either side, running into 
a brown splashed area on the margins of the thorax. 

Maxillae dark shining brown with thick tomentum of fine 
white hairs interspersed with darker brown ones. 

Lip and maxillae reddish-brown, sternum and coxae 
darker with fine clothing of black hairs. 

Abdomen above, dirty white with four discrete broad 
greyish-black bands interspersed with small spots and a similar 
densely-spotted area at sides of abdomen. 

Below, yellowish-white with a broad central black band 
narrowing towards spinnerets, which are likewise black. 

The whole abdomen is clothed with a fine white 
tomentum. 

The eye area, of usual shape, appears white from the 
thickness of the tomentum. 

Legs dark brown, under-surface of tibiae densely clothed 
with white hairs, showing marked contrast with the remain- 
ing joints. 

Total length, 73 mm.; thorax abd., 27 mm. 

This robust Lycosa was found at Whyte-Yarcowie, South 
Australia. 

Type in South Australian Museum. Male unknown. 


DESCRIPTION OF PLATE V. 


Tycosa skeeti, n. sp. 
Nat. size. 
Tycosa perinflata, n. sp. 
Nat. size. 


85 


THE PARASITES OF AUSTRALIAN BIRDS. 


By J. Burton Cietanp, M.D. 
[Read May 11, 1922.] 


Apart from the interest centred in themselves zoologically 
as species and genera of animals, the parasites of birds may 
claim special attention from ornithologists on several grounds 
connected with their hosts. It is with the bird aspect that 
this contribution deals. 

The ecto- and endo-parasites of birds may affect their 
health. In a state of Nature we have little evidence of this 
as far as our native species are concerned. Attention may be 
called, however, to the helminth ova found in tumours in 
the intestine of a black duck. 

In certain cases, ornithologists may perhaps gain con- 
siderable help from a study of bird parasites in establishing 
generic affinities in otherwise doubtful cases. With some 
exceptions, and excluding occasional accidental infections of 
birds of other genera, both such external parasites as 
mallophaga and such internal ones as the helminths are 
probably remarkably specific as regards their hosts: that is, 
are confined to one species of bird only or to a few closely- 
allied species. In a broader sense, this may also apply to 
genera. The reason for this specification is clear. The 
ancestors of the parasites undoubtedly began their parasitic 
career as accidental infestations, individuals gaining access 
to their hosts in some way, being able to resist the efforts 
of these hosts to dislodge them, and being capable of nourish- 
ing and reproducing themselves in their new environment. 
In the course of time they became structurally more and more 
modified to fit themselves for the parasitic life. Modifica- 
tions suitable for one host might be unsuitable for others. 
Passage would more easily be achieved from one host to 
another of the same species. During the period in which 
the parasites were undergoing these marked evolutionary 
changes, their hosts would also be doing the same. Some- 
times the parasites would change structurally more quickly 
or more markedly than their hosts, and then we would have 
perhaps two or more closely-allied species of parasite in one 
specific host or in two or in several closely-related hosts. In 
other cases the parasites might remain more or less 
stationary, whilst considerable structural changes might occur 
in several directions in the descendants of the original host. 


86 


We then might have identical or closely-allied parasites in 
two or more related species or in closely connected genera. 
A generic or even family relationship might thus be shown, 
and it is possible that a disputed point might be settled in 
this way. Thus, supposing a genus X appears to be related 
either to the genus Y or to the genus Z, if X and Y have 
closely related parasites and those of Z are distantly 
connected, then support is found for the relationship with 
Y rather than Z. 

L. Harrison (Parasit., vili., 1915, pp. 88-100), in an 
article on ‘““Mallophaga from Apteryx, and their significance, 
with a note on the Genus fallicola.’”’ deals in an interesting 
way with the vaiue of these ecto-parasites as showing probable 
affinities amongst their hosts. | 

The following extract from Nature (No. 2330, vol. 93, 
June 25, 1914, p. 439) shows another interesting phase of 
this subject:—“From a paper by Mr. H. Victor Jones in 
the February number of the Zoologist on certain parasites 
of birds, we learn that while rooks and the diurnal birds-of- 
prey—probably owing to the strength of their gastric juices— 
are practically free from intestinal infestations of this kind, 
curlews show, on the average, no fewer than 49°5 per head. 
As there seems to be a connection in many species between 
the numbers of external and internal parasites, it is 
suggested that some of the former may serve as hosts for 
the latter during the earlier stages of their development.”’ 

There is clearly very much work still to be done in the > 
parasitology of Australian birds. I have now collected a 
considerable number of mallophaga and worms which await 
description when our few investigators in these subjects have 
time to consider them. Cestodes have already been described 
from 44 species of our birds. J record their occurrence in 
59 species, of which 50 are new hosts. Cestodes, it will be 
seen from the attached list, are rare in our wild parrots, 
have not yet been met with in our cuckoos, are common in 
the honey-eaters, and occur in several of the Ptilonorhyn- 
chide, as well as in other genera. J have not found any in 
the Acanthizas (9 species and 30 individuals) or in Sericorms 
(4 species and 15 individuals). Here it may be mentioned 
that, once helminth parasites are “dropped’’ by a_host- 
species or host-genus, with rare exceptions it is very 
unlikely that such species or genus will ever again become 
infested by such parasites. In other words, these parasites 
are usually so highly specialized that they can only 
exceptionally adapt themselves to hosts of a quite different 
kind, and new true parasites derived from semi-parasites only 
very rarely arise. 


87 


The following summarises the results of previous records 
and my findings in birds examined :— 


Cestodes recorded previously in 45 species. Found by me in 
59, of which 50 are new hosts. 

Adult nematodes recorded previously in 21 species. Found 
by me in 22, of which 15 are new hosts. 

Microfilariae recorded previously in 34 species. | 

Acanthocephala recorded previously in 21 species. Found 
by me in 10, of which 6 are new hosts. 

Trematodes recorded previously in 38 species. Found by me 
ry ft. 

Fleas recorded previously in 1 species. Found by me in 2, 
of which 1 is a new host. 

Hippoboscidae recorded previously in 3 species. Found by 
me in 1, which is a new host. 

Mallophaga recorded previously in 64 species. Found by me 

... _ in 65, of which 54 are new hosts. 

Ticks recorded previously in 1 species. Found by me in 4, 
of which 3 are new hosts. 

Mites recorded previously in 18 species. Found by me in 22, 
of which 21 are new hosts. 

Haemosporidia recorded previously in 47 species. 

Haemoflagellates recorded previously in 12 species. 

In 302 individuals, comprising 132 species, no entozoa 
were detected. Though it is probable that, in some of these, 
parasites were overlooked (and in some of the species they 
have been previously recorded), the number of infested birds 
thus missed is probably small. No ectozoa were detected on 


61 individuals belonging to 46 species. 


The numbers following the names of the birds are those 
of the check-list published in the Emu, vol. xii., 1912-3. 


Pr. I.—ReEcorpDED PARASITES oF AUSTRALIAN BrRDs. 


1. Cestodes.") 


Order CASUARITFORMES. 
Dromaius novae-hollandiae (No. 1).—Davainea australis, Krabbe; 
Cotugnia collini, Fuhr. 
Casuarius australis (No. 4.)—Cestodes in intestine, N.Q., Mac- 
gillivray (Emu, xvii., 1917, p. 80). 
Order COLUMBIFORMES. 
Leucosarcia picata (No. 40).—Davainea sp., Justn. 


Order PODICIPEDIFORMES. 
Podiceps gularis (No. 57).—Taenia novae-hollandiae, Krefft; 
Taenia paradoxa, Krefft. 


(1) Unless a full reference is given, the references to the various 
records will be found in a paper by Dr. T. Harvey Johnston on 
‘Internal Parasites recorded from Australian Birds’? (Emu, vol. 
xli., 1912, p. 105), which forms the basis for this List. 


88 


Order PROCELLARIIFORMES. 


Diomedea exulans (No. 94).—Tetrabothrius sp., Jnstn. 
Diomedea melanophrys (No. 95).—Tetrabothrius sp., Jnstn. 


Order CHARADRITFORMES. 


Lobivanellus lobatus (No. 128).—Gyrocoelia sp., Jnstn. (Ann, Trop. 
Med. and Paras., vili., 1914, p. 108). 

Zonifer pectoralis (No. 130).—Choanotaenia zoniferae, Jnstn. 

Himantopus leucocephalus (No. 142).—Gryocoelia australiensis, 
Jnstn, (Tiaenia coronata, Krefft); Acoleus hedleyi, Jnstn. 
(Taenia rugosa, Krefft); Davainea himantopodis, Jnstn.; 
Hymenolepis sp., Jnstn. 

Gallinago australis (No. 166).—Aploparaksis australis, Jnstn. 

Oedicnemus grallarius (No. 171).—Angularia australis, Maplestone 
(Ann. Trop. Med. and Parasit., xv., No. 4, 1921). 


Order ARDEIFORMES. 
Platalea regia (No. 178).—Cylorchida omalancristrota, Wedl. 
Platibis flavipes (No. 179).—Hymenolepis ibidis, Jnstn. 
Xenorhynchus asiaticus (No. 180).—Clelandia parva, Jnstn. 
Herodias syrmatophorus Settee (No. 184).—Anomotaenia 
asymmetrica, Jnstn.; Bancroftiella glandularis, Fuhrm. 
Noto cakes novae-hollandiae (No. 185).—Bancroftiella glandularis, 
“uhrm. 
Nycticorax caledonicus (No. 191).—Bancroftiella ardeae, Jnstn. ; 
Hymenolepis sp., Jnstn. 


Order ANSERIFORMES. 


Anseranas melanoleuca (No. 199).—Hymenolepis megalops, Nitzsch ; 
Hymenolepis terraereginae, Jnstn. 

Dendrocygna arcuata (No. 204).—Diploposthe laevis, Bl.; Diorchis 
flavescens (Krefft), Maplestone (Ann. Trop. Med. and Parasit., 
xv., No. 4, 1921, p. 403). . 

Anas_superciliosa (No. 208).—Hymenolepis megalops, Nitzsch 
(Taenia cylindrica, Krefft); H. collaris, Batsch. (Taenia bairdii, 
Krefft); Uymenolepis sp., Jnstn.; Diorchis flavescens, 
(Krefft); Fimbriaria fasciolaris, Pall. (Taenia pediformis, 
(Krejft); Diploposthe laevis, Bl. 

Nettium castaneum (No. 209).—Diorchis flavescens (Krefft); 
Diploposthe laevis, Bl.; Hymenolepis collaris, Bat.; Hymeno- 
lepis megalops, Nitzsch (Taenia cylindrica, ‘Krefft); Fimbriaria 
fasciolaris, Pall. 

Spatula rhynchotis (No. 213).—Diorchis flavescens (Krefft). 
Nyroca australis (No. 216).—Diploposthe laevis, Bloch (Taenia 
tuberculata, Krefft;; Diorchis flavescens (Krefft ). 

Biziura lobata (No. 218).—Taenia moschata, Krefft. 


Order PELECANIFORMES. 
Tachypetes (Fregata) aquila (No. 229).—Tetrabothrius sp., Jnstn. 


Order ACCIPITRIFORMES. 
Accipiter torquatus (No. 240).—Anomotaenia accipitris, Jnstn. 


Order PSITTACIFORMES. 
Trichoglossus swainsoni (No. 274).—Moniezia trichoglossi, Linstow. 
Cacatua galerita (No. 291).—Davainea cacatuina, Jnstn. 
Platycercus eximius (No. 311).—Dilepis bancrofti, Jnstn. 


89 


; Order CORACIIFORMES. 
Dacelo gigas (No. 345).—Similuncinus dacelonis, Jnstn. 


Order PASSERIFORMES. 


Petroica goodenovii (No. 394).—Hymenolepis sp., Jnstn. 

Pachycephala rufiventris (No. 430). ions ye punctata, 
Jnstn. (Proc. Roy. Soc. Qland, xxvi., 1914 76). 

Malurus cyaneus (No. 5380). —Choanotaenia Piar Instn, 

Zosterops dorsalis (No. 599).—Zosteropicola clelandi, Jnstn, 

Conopophila albogularis (No. 634)—Davainea conopophilae, Jnstn. 

ee chrysotis (No. 644).—Choanotaenia meliphagidarum, 

nstn, 

Ptilotis leucotis (No. 651).—Choanotaenia eel ghasaearur Instn. 

Meliornis novae-hollandiae (No. 668).—Choanotaenia meliphagi- 
darum, Jnsin. 

Meliornis sericea (No. 669).—Choanotaenia meliphagidarum, Jnstn- 

Entomyza cyanotis (No. 680).—Davainea conopophilae, Instn. 

‘Philemon citreogularis (No. 685).—Davainea conopophilae, Instn. 

Sphecotheres maxillaris (No. 714). —Davainea sphecotheridis, 
Jnstn. (Ann. Trop. Med. and Parasit., vili., 1914, p. 106). 

Chlamydera maculata (No. 722). —Choanotaenia chlamyderae, 
(Krefft ). 

Ptiloris alberti No. 730).—Biuterina clavulus, Linst. 

Corvus coronoides (No. 732).—Davainea sp., Jnstn. 


2. Nematodes (omitting Microfilariae). (2) 


Order GALLIFORMES. 


Catheturus lathami (No. a —Heterakis bancrofti, Jnstn.; Hete- 
rakis catheturinus, Jnstn, 


Order PROCELLARITFORMES. 
Daption capensis (No. 86).—Rictularia shipleyi, Stoss (probably). 


Order ANSERIFORMES. 
Chlamydochen jubata (No. 203).—Heterakis chenonettae, Jnstn. 


Order PELECANIFORMES. 


Phalacrocorax carbo (No. 219).—Ascaris sp., Jnstn. 

Phalacrocorax sulcirostris (No. 220).—Ascaris spiculigera, Rud. 

Plotus novae-hollandiae (No. 224).—Ascaris spiculigera, Rud. 
(Ascaris sp., Krefft). 

Pelecanus conspicillatus (No. 233).—Asearis spiculigera, Rud.(? ) 


Order ACCIPITRIFORMES. 


Falco lunulatus (No. 258).—Filaria sp., Jnstn. 
Hieracidia berigora (No. 259). —Filaria guttata, Schneider. 


Order STRIGIFORMES. 


Ninox boobook (No. 263).—Filaria sp., Jnstn. 3 
Ninox ocellata (No. 264).—Filaria sp., Jnstn. 


(2) For references, vide footnote to List of Recorded Cestodes 
of Australian birds. 
D 


| 
| 
| 
| 
| 
| 
| 


90 


Order CORACIIFORMES. 
Sub-order Poparet. 
Podargus Pe Bancroft (Proc. Roy. Soc. Q’land, 


Se p 
Sub-order HALcyongEs. 


Dacelo leachi (No. 346).—(Filaria) dacelonis, Breinl (Austr. Inst. 
Trop. Med., Rep. for 1911, p. 42). 


Order PASSERIFORMES. 


Flam. CAMPOPHAGIDAE. 
Graucalus melanops (No. 457).—Filaria sp., Jnstn. 


Fam. ME&LIPHAGIDAE. 
Myzantha garrula (No. 672).—Filaria sp., Bancroft. 
Anthochaera_carunculata (No. 675).—Oxyspirura anthochaerae, 
Jnstn. (Proc. Roy. Soc. Q’land, xxiv., 1912, p. 80) (Ceratospira 
anthochaerae, Jnstn.; Ascaris sp., Krefft). 
Annellobia mellivora (No. 677) (recorded as A. lunulata).—Filaria 
sp., Bancroft. 
Acanthogenys rufigularis (No. 679).—Filaria sp., Jnstn. 
Philemon citreogularis (No. 685).—Filaria sp., Jnstn. 
' Fam. CorvipDAe. 
Corvus australis (No. 734).—Fuilaria sp., Bancroft. 


Fam. STREPERIDAE. 


Cracticus destructor (No. 745).—Filaria sp., Bancroft. 
Gymnorhina tibicen (No. 747).—Filaria clelandi, Jnstn. 


2a. Microfilariae. © 


Microfilariae have been described or merely recorded 
from the following species of Australian birds :— 


Order PELECANIFORMES. 


Phalacrocorax sulcirostris (No. 220); P. melanoleucus (No, 228); 
Plotus novae-hollandiae (No. 224) 


Order ACCIPITRIFORMES. 
Accipiter torquatus (No. 240). 


Order PSITTACIFORMES. 

Trichoglossus swainsoni (No. 274), Bancroft (Proc. Roy. Soc. 
Q’land, vi., 1889 [1890]); Glossopsitta pusilla (No. 280) 
Order CORACIIFORMES. 

Podargus strigoides (No. 337); Eurystomus pacificus (No. 341). 


Order PASSERIFORMES. 
Fam, Pirripar£. 


Pitta strepitans (No. 377), Breinl (Austr. Inst. Trop. Med., Rep. 
for 1911, p. 43). 


— 


(3) For references, vide footnote to List of Recorded Cestodes 
of Australian Birds. 


91 


Fam. MUSCICAPIDAE. 
Myiagra planer (No. 444). 


Fam. TIMELIIDAE. 


Psophodes crepitans (No. 476); Pomatorhinus temporalis (No. 478) 
(Pomatostomus frivolus). 


Fam. TURDIDAE. 
Oreocincla lunulata (No. 488). 


Fam, ARTAMIDABR. 


Artamus leucogaster (No. 559); A. sordidus (No. 564) (Artamus 
tenebrosus). 
Fam. PRIONOPIDAE. 


Colluricincla harmonica (No. 566), Cleland (these Trans., xxxix., 
1915, p. 38). 
Fam. (?) 


Struthidea cinerea (No. 576); Corcorax melanorhampus (No. 577). 


Fam. DIcaEIDAE. 
Pardalotus melanocephalus (No. 609), Cleland and Jnstn, (Jour. 
Proc. Roy. Soc. N.S. Wales, xlv., 1911, p. 438). 
Fam. MELIPHAGIDAE. 


- Plectorhyncha lanceolata (No. 621); Myzomela sanguineolenta 
(No. 622); Stigmatops ocularis (No. 639); Ptilotis fusca (No. 
643); Myzantha garrula (No. 672); Anellobia mellivora (lunu- 
lata of Bancroft’s record) (No. 677 7) : Entomyza cyanotis (No. 
680); Philemon citreogularis (No. 685). 


Fam. ORiIoLipAe. 


Oriolus viridis (No. 712). 

Fam. DiIcRURIDAE£. 
Chibia bracteata (No. 716). 

Fam. PTinoNORHYNCIDAE. 
Sericulus chrysocephalus (No. 726). 
Fam. CorvipDAE. 

Corvus coronoides, Corvus australis (Nos. 732, 734). 

Fam. STREPERIDAE. 


pape graculina (No. 735), Bancroft (Proc. Roy. ae Q’land, 
, 1889 [1890] ; Cracticus nigrogularis (No. 741); C. destructor 
(No. 745). 
Gymnorhina tibicen (No. 747).—Microfilaria gymnorhinae, Gilruth, 
Sweet, and Dodd. 


3. Acanthocephala.” 
Order GALLIFORMES. 
Catheturus lathami (No. 7).—Echinorhynchus sp., Jnstn. 


Order CHARADRIIFORMES. 
Numenius cyanopus (No. 145).—Echinorhynchus sp., Jnstn. 


(4) For references, vide footnote to List of Recorded Cestodes 


of Australian Birds. 
D2 


92 


Order ACCIPITRIFORMES. 


Astur cinereus (No. 236).—Centrorhynchus asturinus, Jnstn. 
(Proc. Roy. Soc. Q’land, xxx., 1918,.p. 216). ; 

Astur novae-hollandiae (No. 237).—Centrorhynchus asturinus, 
Jnstn. (loc. cit.). 


ie approximans (No. 238) (A. fasciatus).—Echinorhynchus sp., 
nstn. 


Baza subcristata (No. 254).—C. asturinus, Jnstn. (loc. cit.). 


Order STRIGIFORMES. 
Ninox boobook (No. 268).+Echinorhynchus=Centrorhynchus sp., 


Instn. 
Order CORACIIFORMES. 
Podargus strigoides (No. 337).—EH. sp., Jnstn. 
Haleyon sanctus (No. 349).—E. sp., Jnstn. 


Order MENURIFORMES. 
Menura superba (No. 374).—E. menurae, Jnstn. 


Order PASSERIFORMES. 
e 


F'iam. MUSCICAPIDAE. 


Pachycephala gilberti (No. 432).—Hchinorhynchus pomatostomi, 
Jnstn. and Clel. (arvae, subcutaneous). 


Fam. TiIMELIIDAE. 


Hylacola pyrrhopygia (No. 474).—E. pomatostomi, Jnstn. and 
Clel. (larvae, subcutaneous). 

Psophodes crepitans (No. 476).—E. sp., Jnstn. 

Pomatorhinus temporalis (No. 478).—E. pomatostomi, as above. 

Pomatorhinus superciliosus (No. 479).—EK. pomatostomi, as above. 

Pomatorhinus rubeculus (No. 481).—E. pomatostomi, as above. 


Fam. TurDIDAE. 
Oreocincla lunulata (No. 488).—Echinorhynchus sp., Jnstn. 


Fam. PRIONOPIDAE. 
Grallina picata (No. 575).—E. sp., Jnstn. 


Fam. Paripae£. 
Aphelocephala leucopsis (No. 578).—E. pomatostomi, as above. 


Fam. CERTHIIDAE. 
Climacteris melanura (No. 589) (C. wellsi)—E. pomatostomi, as 
above. 
Fam. MELIPHAGIDAE. 


‘Meliornis novae-hollandiae (No. 668).—E. sp., Justn. 


4. Trematodes. © 


Order COLUMBIFORMES. 


Leucosarcia picata (No. 40).—Harmostomum pulchellum, 8S. J. 
Johnston (N.S. Wales). 


(5) Compiled from S. J. Johnston’s article ‘‘On the Trematodes 
of Australian Birds’? in Jour. Proc. Roy. Soc. of N.S. Wales, 1., 
19lb peeled 4 = ; 


93 


Order RALLIFORMES. 
Porphyrio melanonotus (No. 55).—Echinostomum hilliferum, Nicoll. 


Order LARIFORMES. 


Sterna cristata (No. 107) (S. bergii).—Lyperosomum megastomum, 
S. J. Johnston (N.S. Wales); Holostomum musculosum, S. J. 
Johnston (N.S. Wales). ‘ 

Larus novae-hollandiae (No. 119).—Austrobilharzia terrigalensis, 

. J. Johnston (N.S. Wales); Holostomum hillii, S. J. 
Johnston (N.S. Wales). 


Order CHARADRIIFORMES. 


Lobivanellus lobatus (No. 128)._Haematotrephus  consimilis, 
Nicoll; Echinostomum ignavum, Nicoll. 

Charadrius fulvus (No. 1382) (C. dominicus).—Acanthoparyphium _ 
spinulosum, S. J. Johnston (N.S. Wales); Levinseniella 
howensis, S. J. Johnston (Lord Howe Island). 

Himantopus leucocephalus (No. 142).—Haematotrephus adelphus, 
S. J. Johnston (S. Austr.). 

Numenius cyanopus (No. 145).—Himasthla harrisoni, S. J. 
Johnston (Q’land). 

Limosa novae-hollandiae.—Cyclocoelum taxorchis, S. J. Johnston 
(Lord Howe Island). 

(Gidienemus grallarius (No. 171) (Burhinus grallarius).—Platy- 
notrema biliosum, Nicoll; P. jecoris, Nicoll. 


Order GRUIFORMES. 


Antigone australasiana (No. 174).—Allopyge antigones, S. J. 
. Johnston; Echinostomum australasianum, Nicoll. 


Order ARDEIFORMES. 


Ibis anes (No. 175).—Patagifer acuminatus, S. J. Johnston 
*Jand). 

Carpe See (No. 176).—Hchinostoma acuticauda, Nicoll 
*land). 

Platalea regia (No. 178).—Orchipedum sufflavum, Nicoll; Patagifer 
bilobus, Rud. 

Plegadis falcinellus (No. 177).—Patagifer bilobus, Rud. 

Xenorhynchus asiaticus (No. 180).—Chaunocephalus ferox, Rud. 

Herodrias (Syrmatophorus) timoriensis (No. 184).—Patagifer 
fraternus, S. J. Johnston (Q’land); Echinoparyphium 
oxyurum, S. J. Johnston (Q’land). 

Notophoyx novae-hollandiae (No. 185) (Ardea novae-hollandiae).— 
Holostomum simplex, S. J. Johnston; H. repens, Chase (Proc. 
Linn. Soc. N.S. Wales, xlv., 1921, p. 500) (N.S. Wales). 

Nycticorax caledonicus (No. 191).—Clinostomum hornum, Nicoll. 


Order ANSERIFORMES. 

Chenopis atrata (No. 198).—Hemistomum intermedium, S. J. 
Johnston; Hyptiasmus magnus, S. J. Johnston (Vict.); 
Notocotylus attenuatus, Rud. 

Anseranas melanoleuca (No. 199) (A. semipalmata).—Typhlocoelum 
reticulare, S. J. Johnston. 

Nettapus pulchellus (No. 200).—Notocotylus attenuatus, Rud. 

Anas superciliosa (No. 208).—Echinostomum revolutum, Foel. ; 
Notocotylus attenuatus, Rud. 


94 


Order PELECANIFORMES. 
Phalacrocorax melanoleucus (No, 223).—Dolichosaccus solecarius, 


S. J. Johnston (N.S. Wales); Echinochasmus tenuicollis, S. J. 
Johnston (N.S. Wales). 


Plotus novae-hollandiae (No. 224).—Clinostomum australiense, 
S. J. Johnston (Q’land). 


Order ACCIPITRIFORMES. 
Haliaetus leucogaster (No. 246).—Scaphanocephalus australis, 
Johnston. 
Hieracidea berigora (No. 259).—Opisthorchis obsequens, Nicoll 
(Q’ land). 
H. orientalis (No. 260).—Echinochasmus prothovitellatus, Nicoll 


(Q’ land). 
Order STRIGIFORMES. 
Ninox boobook. (No. 268).—Lyperosomum harrisoni, S. J. Johnston 
(N.S. Wales); Strigea promiscua, Nicoll (Q’land). 
N. maculata (No. 265).—Strigea promiscua, Nicoll (Q’land); 
Hemistomum brachyurum, Nicoll (Q’land); H. triangulare, 
S. J. Johnston (N.S. Wales). 


Order CORACITFORMES. 
Podargus strigoides (No. 337).—Echinostomum elongatum, Nicoll. 
Dacelo gigas (No. 345).—Hemistomum triangulare, S. J. Johnston 
(N.S. Wales); Strigea flosculus, Nicoll. 


Order COCCYGES. 
Centropus phasianus (No. 373).—Echinostomum emollitum, Nicoll. 


Order PASSERIFORMES. 
Petrochelidon ariel (No. 387).—Plagiorchis clelandi, S. J. Johnston 
(N.S. Wales). 
Microeca fascinans (No. 388).—Echinoparyphium harveyanum, 
S. J. Johnston (Q’land). 


Saar ee (No. 687).—Plagiorchis spatulatus, S. J. Johnston 

land). 

Chibia bracteata (No. 716).—Plagiorchis (Lepoderma) nisbettii, 
Nicoll; -Prosthogonimus vitellatus, Nicoll. 

Strepera arguta (No. 786) (S. versicolor).—Lyperosomum parvum, 
Re J. Johnston (N.S. Wales). 


5. Siphonaptera (Fleas). 


Order SPHENISCIFORMES. 


Eudyptula minor (No. 62).—Parapsyllus australiacus, Roths. (Nov. 
Zool., xvi., 1909, p. 62) (P. longicornis, Jord. and Roths. 
[nec. Enderl., err. determ.)). 


6. Diptera. 
Order ACCIPITRIFORMES. 


White Hawk.—Ornithoctona nigricans, Leach (S. Q’land) (vide 
W. W. Froggatt in ‘‘Australian Insects’’). 


Order STRIGIFORMES. 
Ninox boobook (No. 263).—Ornithomyia perfuga, Spajer, on an 
owl, probably this species, near Brisbane (vide Froggatt, loc. 
cit. ). 


a 


95 


Order MENURIFORMES. 


| Menura superba (No. 374).—Larvae of a Muscid fly subcutaneously 


(Gubert, Emu, xix., 1919, p. 48). 


Order PASSERIFORMES. 


Stipiturus malachurus (No. 545).—Ornithomyia stipituri, Schiner 
(Zool. Voy. Novara, 1850) (vide Froggatt, loc. cit.). 

Glyciphila fulvifrons (No. 629).—Larvae of a Muscid fly sub- 
cutaneously (Gilbert, loc. cit., p. 49). 

Meliornis novae-hollandiae (No. 668).—Larvae of a Muscid fly sub- 
cutaneously (Gilbert, loc. cit., p. 49). 

Meliornis sericea (No. 669).—Larvae of a Muscid fly subcutaneously 
(Gilbert, loc. cit., p. 48). 

Anthus australis (No. 687).—Fly larvae attached to body (Harvey, 
Emu, xix., 1919, p. 40; Q’land). 

Mr. Froggatt also states that Hippoboscid flies occur on our 
fruit-pigeons, swallows, and fly-catchers. 


7. Mailophaga. °° 
Order CASUARIIFORMES. 


Dromaius novae-hollandiae (No. 1).—Degeeriella asymmetrica 
(Nitzsch) (syn. Nirmus setosus, Le Souéf and Bullen) (Q’land, 
N.S. Wales, Vict.). 


Order GALLIFORMES. 


Catheturus lathami (No. 7).—Goniocotes fissus, Rud. (from Tale- 
gallus lathami); G. macrocephalus, Taschenb. (from T. 
lathami); Japeurus ischnocephalus, Taschenb. (from T. 
lathami); L. crassus, Rud. 

Synoicus australis (No. 9).—Goniodes elongatus, Piaget (Vict.); 
G. retractus, Le Souéf (Vict.). 

Excalfactoria australis (No. .10).—Lipeurus acuminatus, Piaget; 
Goniodes elongatus, Piaget (syn. G. longus, Le Souéf); 
Menopon pallipes, Piaget. 


Order COLUMBIFORMES. 


Megaloprepia magnifica (No. 21).—EHsthiopterum columbae (L.) 
(syn. Lipeurus baculus, Nitzsch, and N. angustus, Rud.) 
(from Carpophaga magnifica). 

Macropygia phasianella (No. 25).—Colpocephalum albidum, Giebel 
(from Columba phasianella). 

Phaps chalcoptera (No. 30).—Goniocotes flavus (Rud.); Esthiop- 
terum solutes (L.) (Tas.); Colpocephalum albidum, Gebel. 

Leucosarcia pictata (No. 40).—Esthiopterum columbae (L.) (from 
Leucosarca plicata). 


(6) These records are compiled almost entirely from Prof. V. 
L. Kellogg’s article ‘‘Mallophaga’’ in Wytman’s ‘‘Genera 
Insectorum,” 1908, from Johnston and Harrison’s ‘‘Census of Aus- 
tralian Mallophaga’’ in Proc. Roy. Soc. Q’land, xxiv., 1912, and 
particularly from Harrison’s ‘‘Census of Mallophaga in Para- 
sitology,’’ ix., 1916, pp. 1-152. Full references are only given for 
species not included in these lists. In many cases, though the 
host occurs in Australia, the parasite has not actually as yet been 
recorded for this country. Australian occurrences are indicated. 


96 


Order RALLIFORMES. 
Rallina tricolor (No. 44).—Rallicola bisetosa (Piaget) (syn. 
Oncophorus bisetosus, Piaget). 
Tribonyx ventralis (No. 52).—Goniodes cornutus, Rud. (straggler ; 
L. H.).; Philopterus flavopunctatus, Rud. 
Porphyrio melanonotus (No. 55).—Rallicola (Oncophorus) « fallax 
(Piaget) (from Porphyrio melanotus, Australia). 


| Order SPHENISCIFORMES. 


. EKudyptula minor (No. 62).—Austrogoniodes waterstoni, Cummings 
(Bull. Ent. Rev., v., 1914, p. 173, f. 8). 


Order PROCELLARIIFORMES. 

Ossifraga gigantea (No. 85).—Hsthiopterum obscurum (Rud.) 
(syn. Lipeurus melanocnemis Gebel) (from Procellaria 
gigantea). - 

Daption capensis (No. &86).—Esthiopterum (Lipeurus) gurlti 
(Taschenb.) (from Procellaria capensis); E. nigrolimbatum 
(Giebel) (syn. KE. mutabile, Piaget); E. fuliginosum, Taschenb. 
(syn. EK. testaceum, Taschenb.); Ancistroma vagelli, Fabr. 
(syn. A. procellariae, Westwood), N.S. Wales. ; 

Diomedea exulans (No. 94).—Docophoroides brevis, Dufour (syns. 
D. dentatus, Giebel, and D. taurus, Nitzsch); Esthiopterum 
pederiforme, Dufour (syns. Docophorus thoracicus, Nitzsch ; 
Nirmus angulicollis, Giebel; and L. breviceps, Piaget); E. 
hyalineum (Neum.); EK. fuliginosum, Taschenb. (syn. Lipeurus 
fuliginosus, Taschenb.); Menopon affine, Piaget. 

Diomedea chlororhynchus (No. 98).—Esthiopterum (Lipeurus) 
fuliginosus, Taschenb. 


Order LARIFORMES. 


Sterna cristata (No. 107).—Colpocephalum crassipes, Piaget (from 
S. poliocera=S. bergii=this species). 


Order CHARADRIIFORMES. 
Tringa canutus (No. 164).—Degeeriella (Nirmus) holopaea 
(Nitzsch). 
Parra gallinacea (No. 168).—Parricola sulcata, Piaget (syn. 
Oncophorus sulcatus, Piaget). 


Order GRUIFORMES. 


Antigone australasiana (No. 174).—Philopterus integer, Nitzsch 
(syn. Docophorus integer, Nitzsch); Philopterus novae- 
hollandiae, Giebel (syn. D. novae-hollandiae, Giebel); 
Esthiopterum (Lipeurus) giganteum (Le Souéf and Bullen) 
Q’land, N.S: Wales, Vict.); E. (Lipeurus) gruis (L.) (syn. L. 
hebraeus, Nitzsch) (Q’land, N.S. Wales, Vict.). 


Order ARDEIFORMES. 


Ibis molucca (No. 175).—Esthiopterum ibidis, Harris. (syn. Lipeurus 
ibis, Le Souéf and Bullen, from Threskiornis strictipennis, 
Australia). 

Platibis flavipes (No. 179).—Ornithobius fuscus, Le Souéf (Pa 
straggler). ° 

Xenorhynchus asiaticus (No. 180).—Philopterus horridus, Gtebel 
(from Ciconia australis). 


97 


Ardea cinerea (No. 182).—Colpocephalum decimfasciatum, Bois. 

and Lacord. (syn. C. importunum, Nitzsch); Esthiopterum 

i ardeae (L.) (syn. Lipeurus leucopygus, Nitzsch). 

Notophoyx novae-hollandiae (No. 185).—Philopterus longipes, 
Rud.; Esthiopterum (Lipeurus) unguiculatum (Piaget) (from 
Herodias novae-hollandiae). 


Order ANSERIFORMES. 

Chenopis atrata (No. 198).—Esthiopterum megacerus, Jnstn. and 
Harris. (syns. Lipeurus anatis megaceros, Jnstn. and Harvis., 
and L. squalidus, Nitzsch, var. attenuata, Piaget); Orni- 
thobius fuscus, Le Souéf (Vict.); Trinoton nigrum, Le Souéf 
(Vict.); Colpocephalum castaneum, Piaget (from Cygnus 
atratus). 

Anseranas melanoleuca (No. 199).—Heteroproctus hilli, Harrison 
(Parasit., vil., 1914-5, p. 394) (Northern Territory). 

Cereopsis novae-hollandiae (No. 202).—Esthiopterum australe 
Rud.) (syn. Lipeurus australis, Rud.) (from Coreopsis novae- 

ollandiae). 

Nettium gibberifrons (No. 210).—Esthiopterum crassicorne 
(Scopoli) (syns. Lipeurus anatis major, Piaget, and L. 
squalidus, Nitzsch, var. major, Piaget (from Anas gibberifrons). 

Nyroca australis (No. 216).—Esthiopterum crassicorne (Scopoli) 
(syn. EK. nyrocae [Rud.)). 


, 


Order PELECANIFORMES. 


Phalacrocorax carbo (No. 219).—Degeeriella (Nirmus) interrupta, 
Piaget; Esthiopterum mergiserrati, Degeer (syn. Lipeurus 
‘temporalis, Nitzsch); E. longicorne (Piaget); E. toxocerum 
(Nitzsch); Menopon brevipalpe, Piaget. 


_ Phalacrocorax sulcirostis (No. 220).—Esthiopterum (Lipeurus) 


setosum (Piaget); E. confuscum, Bag. and Hall (syn. E. 
brevicorne, Piaget); E. acutifrons (Rud.) (syn. E. dispar, 
Piaget); Menopon subrotundum, Piaget. 

Sula australis (No. 225).—Philopterus (Docophorus) brevianten- 
natus (Piaget); Eschiopterum (Pectinopygus, Lipeurus) 
gyricornis (Denny); Menopon albescens, Piaget. 


Order ACCIPITRIFORMES. 


Haliaetus leucogaster (No. 246).—Colpocephalum  flavescens, 
Nitzsch. . 

Order PSITTACIFORMES. ; 

Trichoglossus swainsoni (No. 274) (T. novae-hollandiae).—EKomeno- 
pon denticulatum, Harrison (Parasit., vii., 1914-5, p. 385) 
(N.S. Wales); Psittaconirmus australis, Harrison (loc. cit., 
p. 403) (N.S. Wales). 

Ptilosclera versicolor (No. 277).—Eomenopon denticulatum, Harri- 
son (loc. cit.) (N.S. Wales). 

Glossopsitta porphyrocephala (No. 279).—Psittaconirmus australis, 
Harrison (loc. cit.) (W. Austr.). ; 

Microglossus aterrimus (No. 283).—Menopon commissum, Newm. ; 
Degeeriella (Nirmus) paraboliceps (Piaget) (from Psittacus 
aterrimus); Colpocephalum temporale, Piaget (from Macro- 
glossus aterrimus). : 

Calyptorhynchus leachi (No. 289).—Esthiopterum (Lipeurus) 
cireumfasciatum (Piaget) (from Colyptorhynchus leachi). 


| 
| 


98 


Cacatua galerita (No. 291).—Hsthiopterum capreolum, Gervais 
(syn. Lipeurus albus, Le Souéf and Bullen) (Australia). 
Cacatua roseicapilla (No. 295).—Degeeriella eos (Rud.) (syns. 

Nirmus eos, Rud., and N. tenuis, Rud,) (from Plictolophus 
(Psittacus) roseocapillus and Cacatua (Psittacus) eos). 
Calopsitta novae-hollandiae (No. 298).—Paragoniocotes fasciatus, 

Piaget (from Nymphicus novae-hollandiae). 
Polytelis barrabandi (No. 299).—Philopterus (Docophorus) angusto- 
clypeatus (Piaget) (from Platycercus barrabandi); P. (D.) 
forficula (Piaget) (from Platycercus barrabandi); Colipo- 
cephalum trimaculatum, Piaget (from Platycercus barrabandi). 
Polytelis melanura (No. 300).—Esthiopterum (Lipeurus) cireum- 
fasclatum (Piaget) (from Platycercus melaneura). 


Aprosmictus scapulatus (No. 303).—Philopterus forficula (Piaget) 


eee ua ated ner forficula, Piaget) (from Platycercus scapu- 

atus). 

Platycercus pennanti (No. 304).—Philopterus (Docophorus) for- 
ficula (Piaget). , 

Platycercus pallidiceps (No. 308).—Colpocephalum trimaculatum, 
Piaget (from Platycercus palliceps). 

Platycercus eximius (No. 311).—Philopterus (Docophorus) forficula 
(Piaget); Menopon pteropsittacus, Harris. (syn. M. psittacus, 
Le Souéf and Bullen) (Australia). 

Barnardius barnardi (No. 315).—Philopterus forficula (Piaget) 
(from Platycercus baueri and P. zonarius). 

Pezoporus formosus (No. 334) (P._ terrestris).—Degeeriella 
divergens, Newm. 


Order CORACIIFORMKHS. 

Dacelo gigas (No. 345).—Philopterus (Docophorus) delphax 
(Nitzsch) (from Dacelo gigantea); Degeeriella -(Nirmus) 
bracteata (Nitzsch) (from Dacelo gigantea); D. (Nirmus) 
goniocotes (Piaget) (Madagascar); Menopon infumatum, 
Piaget (Madagascar). 


Order COCCYGES. 


Cacomantis flabelliformis (No. 362).—Philopterus (Docophorus) 
laticlypeatus (Piaget) (from Cuculus flabelliformis, New 
Holland). 

Scythrops novae-hollandiae (No. 372).—Philopterus acutus, Rud. ; 
P. (Docophorus) obcordatus (Piaget); Degeeriella lipeuriformis 
Rud.) (syns. Nirmus lipeuriformis, Rud.; N. chelurus, 

itzsch); Myrsidea (Menopon) platygaster (Giebel). 


Order MENURIFORMES. 


Menura superba (No. 374).—Degeeriella menuraelyrae (Coinde) 
(syns. Philopterus (Docophorus) paraboliceps (Piaget); Nirmus 
submarginellus, Nitzsch; N. submarginalis, Burm.; and N. 
menura, Le Souéf and Bullen). 

Johnston and Harrison consider Kellogg’s record of Degeeriella 
(Nirmus) marginalis, Nitzsch, as an error. : 

Menura victoriae (No. 375).—Esthiopterum menura, Le Souéf and 
Bullen (syn. Lipeurus menura, Le Souéf and Bullen) (Vict.); 
Menopon menura, Le Souéf and Bullen (Vict.); Degeeriella 
menuraelyrae (Coinde) (syns. see above) (Vict.). 


99 


Order PASSERIFORMES. 


Fam. DiIcarIDAE. 
Pardalotus punctatus (No. 606).—Menopon sp., Giebel. 


Fam. MELIPHAGIDAE. 
Glyciphila fasciata (No. 631).—Goniocotes candidus, var. pellucidus, 
Piaget (probably a straggler; J. and H). 
Tropidorhynchus corniculatus (No. 684).—Homenopon denticulatus, 
Harrison (Parasitol., vii., 1914-5, p. 385) (N.S. Wales; straggler 
on this host). 


Fam. PLoceIpDAr. 


Poephila gouldiae (No. 709) (P. mirabilis)—Machaerilaemus lati- 
frons, Harrison (Parasitol., vii., 1914-5, p. 390). 


Fam. PT1noNORHYNCHIDAE. 

Ptilonorhynchus holosericeus (No. 718).—Menopon ptilonorhynchi, 
Ponton; Philopterus ptilonorhynchi, Ponton (syn. Docophorus 
erandiceps (Nitzsch) (from Ptilonorhynchus holosericeus) ; 
Degeeriella pontoni, Jnstn. and Harrison (syn. Nirmus 
nitzschi, Ponton) (from Ptilonorhynchus holosericeus). 

Sericulus chrysocephalus (No. 726).—Degeeriella (Nirmus) hectica 
(Nitzsch). 

Fam. CorvIpDAE. 
Strepera graculina (No. 735).—Colpocephalum vinculum, Le Souéf 
- and Bullen (Australia). 

Gymnorhina. tibicen (No. 747).—Degeeriella bimaculata (Piaget) 
(syn. Nirmus bimaculatus, from Baryta tibicen). 

Gymnorhina leuconota (No: 750).—Degeeriella semiannulata 
(Piaget), (syn. Nirmus semiannulatus, Piaget, from Baryta 
leuconota); Degeeriella (Nirmus) varia, Nitzsch (probably a 
stray, Rotterdam). 

8. Acarina. 
(a) Super-family IXODOIDEA. | 

Host probably birds (marine).—Ixodes tasmani, Newm. Collected 
by Verreaux, the ornithologist (1847) in Tasmania (vide Nutt. 
and Warb., Ticks, pt. i1., 1911, p. 245). 

Host marine birds.—Ixodes putus (Pickh.-Cambridge). Recorded 
by Neumann from King Island (Tas. ?) (vide Nutt. and Warb., 
Ticks, pt. ii., 1911, p. 261). 


Order PASSERIFORMES. 
Fam. HirvuNDINIDAE. 
Petrochelidon ariel (No. 387) (Lagenoplastes  ariel).—Argas 
lagenoplastes, Frogg. (Proc. Linn. Soc. N.S. Wales, 1906, 
p. 408). Recorded for Merriwa and Narromine, N.S. Wales, 
and for Q’land. 


(b) Super-family ORIBATOIDEA. 
_ Fam. Anaue@eEsipAr (‘‘Bird Mites’’). 
[For the following records, I am indebted to Mr. W. J. Rain- 
bow’s ‘‘A Synopsis of Australian Acarina’’ (Rec. of Austr. Mus., 
vol. vi., pt. 3, p. 181), where the full references will be found. ] 


Order CHARADRIIFORMES. 
Lobivanellus lobatus (No. 128).—Trouessartia caudacuta, Troues. 


100 


Order ARDEIFORMES. 


Ibis molucca (No. 175).—Freyana (Eufreyana) tarandus, Jrowes 
et Neuwm.; Alloptes corymbophorus, Troues et Neum. 


Order ACCIPITRIFORMES. 
Haliastur leucosternus (No, 247) (H. indicus, var. girrenera).— 
Pterolichus (Kupterolichus) phylloproctus, var. minor. Mégn. 
et Troues; P. (Pseudalloptes) aquilinus, var. milvulina, 


Troues. 
‘ Order PSITTACIFORMES. 

Trichoglossus swainsoni (No. 274) (T. novae-hollandiae).—Ptero- 
lichus (Protolichus) brachiatus, var. crassior, Troues. 

Glossopsitta concinna (No. 278). —Pterolichus (Protolichus) brachi- 
ae var. crassior, Troues; P. (Protolichus) falculiger, Troues; 

(Pseudalloptes) cultriventris, Troues. 

eae aterrimus (No. 283). —Pterolichus (Protolichus) 
favettei, Trowes. 

Caly torhynchus macrorhynchus (No. 287).—Pterolichus (Pseudal- 
(a) spathuliger, Trowes. 

Platycereus pennanti (No. 304).—Pterolichus (Protolichus) chira- 
gricus, Mégn. et Troues; Protalges cartus, Troues. 

Platycercus flaveolus (No. 306), —Pterolichus (Protolichus) chira- 
gricus, Mégn. et Trowes; P. (Protolichus) veliger, Mégn. et 


Psephotus xanthorrhous (No. 319a).—Pterolichus (Protolichus). 
favettei, T'roues. 

Psephotus haematonotus (No. 324).—Analges tetracentrus, Trowes. 

Melopsittacus undulatus (No. 333).—Pterolichus (Protolichus) 
lunula, Robin. 

Pezoporus ’formosus (No. 334) (P. terrestris):—Pterolichus (Proto- 
lichus) chiragricus, Mégn. et Troues. 


Order MENURIFORMES. 
Menura superba (No. 374).—Alloptes major, Trouwes. 


Order PASSERIFORMES. 
Fam. DIcAEIDAE. 
Dicaeum hirundinaecum (No. 602).—Alloptes securiger, Troues. 
Fam. MELIPHAGIDAE. 

Glycyphila fasciata (No. 631).—Protalges australis, Troues; 

Pterodectes manicatus, Trowes. 
Meliornis sericea (No. 669).—Alloptes lobulatus, Trowes. 

7 Fam. PT1noNORHYNCHIDAE. 


Sericulus chrysocephalus (No. 726) (S. melinus).—Pterodectes 
paradisiacus, 7J'rowes. 


9. Haematozoa ”) 
(a) HAEMOSPORIDIA. 
Order GALLIFORMES. 
Catheturus lathami (No. 7).—Halteridium sp. 


(7) For references, vide footnote to List of Recorded Cestodes 
of Australian Birds. 


101 


Order COLUMBIFORMES. 
Lamprotreron superba (No. 20).—Haemoproteus (Halteridium) 


columbae (No. 20), Celli et Fel., Breinl (Austr. Inst. Trop. 
Med., Rep. for 1911, p. 38). 


Order ARDEIFORMES. 


Notophoyx novae-hollandiae (No. 185).—Haemoproteus (Hal- 
teridium danilewskyi, Grassi et Fel., Breinl (Austr. Inst. 
Trop. Med., Rep. for 1911, p. 38). 


Order ANSERIFORMES. 
Chenopis atrata (No. 198).—Proteosoma biziurae, Gilr., Sweet et 
Dodd, ? Clel. (Trans. Roy. Soc. S. Austr., xxxix., 1915, p. 27)- 
Nettium castaneum (No. 209).—Halteridium sp. 
Biziura lobata (No. 218).—Proteosoma biziurae. 


Order ACCIPITRIFORMES. 


Haliastur leucosternus (No. 247) (H. girrenera).—Haemoproteus 
(Halteridium) danilewskyi, Grassi et Fel., Breinl (Austr. Inst. 
Trop. Med., Rep. for 1911, p. 38). 

Faleo hypoleucos (No. 256).—Plasmodium (Proteosoma) praecox, 
Grassi et Fel., Breinl (Austr. Inst. Trop. Med., Rep. for 1911, 


p. 34). 
Order STRIGIFORMES. 


Ninox boobook (No. 263).—Halteridium sp.; Haemoproteus (Hal- 
teridium) noctuae, Celli et Fel., Brewnl (Austr. Inst. Trop. 
Med., Rep. for 1911, p. 37). 

Ninox strenua (No. 268).—Halteridium sp., Clel. (Trans. Roy. Soc. 
S. Austr., xxxix., 1915, p. 29). 


Order PSITTACIFORMES. 
Platycercus adelaidae (No. 305).—Halteridium sp. 


Order CORACIIFORMES. 
Sub-order Poparet. 
Podargus strigoides (No. 337).—Leucocytozoon sp., Clel. (Trans. 
Roy. Soc. S. Austr., xxxix., 1915, p. 30). 
Sub-order Hatcyones. 
_ Dacelo gigas (No. 345).—Halteridium sp. 
Sub-order MeERopss. 
Merops ornatus (No. 352).—Halteridium sp. 


Order COCCYGES. 


Eudynamis cyanocephala (No. 371).—Haemoproteus danilewskyi, 
Grassi et Fel, Breinl (loc. cit.). 


Order PASSERIFORMES. 
Fam. MUSCICAPIDAE. 


Microeca fascinans (No. 388).—Halteridium sp. 

Petroica phoenicea (No. 393).—Halteridium sp. 

-Gerygone albogularis (No. 402).—Halteridium sp., Clel. (Trans. 
Roy. Soc. S. Austr., vol. xxxix., 1915, p. 29). 

Myiagra nitida (No. 446).—Halteridium sp. 


102 


Fam. TIMELIIDAE. 
Pomatorhinus superciliosus (No. 479).—Halteridium sp. 


Fam. TurpDIDAE. 
Oreocincla lunulata (No, 488).—Halteridium sp. 


Fam. SYLVIIDAE. 


Megalurus gramineus (No. 496).—Haemoproteus danilewskyi, 
Grassi et Fel., Breinl (loc. cit.). 


Fam. P 


Grallina picata (No. 575).—Halteridium sp. 
Corcorax melanorhamphus (No. 577).—lLeucocytozoon anellobiae. 


Fam. PARIDAE. 
Aphelocephala leucopsis (No. 578).—Halteridium sp. 


Fam. ZOSTEROPIDAE. 
Zosterops dorsalis (No. 599).—Halteridium sp. 


Fam. DICAEIDAE. 


Dicaeum hirundinaceum (No. 602).—Halteridium sp., Clel. (Trans. 
Roy. Soc. S. Austr., xxxix., 1915, p. 29). 

Pardalotus melanocephalus (No. 609).—Halteridium sp., Clel. and 
Ae te (Jour. and Proc. Roy. Soc. N.S. Wales, xlv., 1911, 
p. 

Fam. MELIPHAGIDAE. 

Melithreptus brevirostris (No. 619).—Halteridium sp. 

Myzomela sanguineolenta (No. 622).—Halteridium sp.; Leucocyto- 

zoon anellobiae. 

Ptilotis fusca (No. 643).—Halteridium sp.; Leucocytozoon 
anellobiae. 

Ptilotis sonora (No. 646).—Halteridium sp. 

Ptilotis chrysops (No. 648).—Halteridium sp. 

Ptilotis plumula (No. 658).—Halteridium sp. . 

Ptilotis penicillata (No. 661).—Halteridium sp., Clel. (Trans. Roy. 
Soc. S. Austr., xxxix., 1915, p. 30). 

Meliornis novae-hollandiae (No. 688).—Halteridium sp. 

Myzantha garrula (No. 672).—Halteridium sp.; Leucocytozoon 
anellobiae. 

Myzantha flavigula (No. 674).—Halteridium sp. 

Anellobia mellivora (No. 677).—Leucocytozoon  anellobiae; 
Haemoproteus (Halteridium) danilewskyi, Grass: et Fel., 
Breinl (Austr. Inst. Trop. Med., Rep. for 1911, p. 38). 

Acanthogenys rufigularis (No. 679).—Halteridium sp., Clel. (Trans. 
Roy. Soc. S. Austr., xxxix., 1915, p. 30); Leucocytozoon sp., 
Clel. (loc. cit., p. 31). 

Entomyza cyanotis (No. 680).—Halteridium sp.; Leucocytozoon 
anellobiae. 

Tropidorhynchus corniculatus (No. 684).—Halteridium = sp.; 
Haemoproteus (Halteridium) danilewskyi, Grassi et Fel., 
Breinl (Austr. Inst. Trop. Med., 1911, p. 37); Leucocytozoon 
sp. (Breinl, etc., p. 37). 


Fam. ORIOLIDAE. 


Oriolus viridis (No. 712).—Halteridium sp.; Leucocytozoon anel- 
lobiae. 
Sphecotheres maxillaris (No. 714).—Leucocytozoon anellobiae. 


103 


Fam. Dicruripaz. 

Chibia bracteata (No. 716).—Haemoproteus (Halteridium) 
danilewskyi, Grassi et Fel., Breinl (Austr. Inst. Trop. Med., 
FOlt, ps 38). 

Fam. PTILONORYHNCHIDAE. 

Chlamydera orientalis (No. 724a).—Haemoproteus (Halteridium) 
danilewskyi, Grassi et Fel., Breinl (Austr. Inst. Trop. Med., 
IEW Se). 


Fam. CorvipDae. 


Cracticus destructor (No. 745).—Haemoproteus (Halteridium) 
danilewskyi, Grassi et Fel., Breinl (loc. cit.). 


(6) HAEMOFLAGELLATES. 
Order ARDEIFORMES. 


Notophoyx novae-hollandiae (No, 185).—Trypanosoma notophoyxis, 
Breil (Austr. Inst. Trop. Med., Rep. for 1911, p. 38). 


Order ACCIPITRIFORMES. 


Haliastur leucosternus (No. 247) (H. girrenera).—Trypanosoma 
avium, Dan. (T. majus, Dan.), Breinl (Austr. Inst. Trop. 
Med., Rep. for 1911, p. 31). 

Falco hypoleucos (No. 256).—Trypanosoma avium, Dan., Breinl 
(Austr. Inst. Trop. Med., Rep. for 1911, p. 31). 


Order STRIGIFORMES. 


Ninox boobook (No. 263).—Trypanosoma sp., Breinl (Austr. Inst. 
Trop. Med., Rep. for 1911, p. 34). 


Order PASSERIFORMES. 


Fam. MUScCICAPIDAE. 
Microeca fascinans (No. 388).—Trypanosoma anellobiae. 


Fam. MELIPHAGIDAE. 
Myzomela sanguineolenta (No. 622).—Trypanosoma sp., Clel. 
(Trans. Roy. Soc. S. Austr., xxxix., 1915, p. 31). 
Ptilotis fusca (No. 643).—Trypanosoma anellobiae. 
Ptilotis chrysops (No. 648).—Trypanosoma sp., Clel. (Trans. Roy. 
Soc. S. Austr., xxxix., 1915, p. 3 
Anellobia mellivora (No. 677) (A. chrysoptera).—Trypanosoma sp. 
Entomyza cyanotis (No. 680).—Trypanosoma sp. 


Fam. ORIOLIDAE. 
Oriolus viridis (No. 712).—Trypanosoma sp. 
Fam. PTILONORHYNCHIDAE. 


Chlamydera orientalis (No. 7244).—Trypanosoma chlamydoderae, 
Breinl (Aust. Inst. Trop. Med., Rep. for 1911, p. 32). 


: VARIOUS PASSING RECORDS. 


Eudyptula minor (No. 62).—Nicholls (Emu, xvii., 1918, pp. 129, 
130) records the following parasites as present in or on four 
birds, viz.:—(1) Round worms in upper part of stomach, fleas ; 
(2) worms, lice, and fleas; (3) small round worms, lice; (4) 
worms, lice. 


104 


Phalacrocorax hypoleucus (No. 222).—In three of twelve birds 
examined in South Australia in March, 1917, parasitic worms 
were noted by Capt. S. A. White (Emu, April; 1918, pp. 214, 
215). Capt. White exhibited two tubes of parasitic worms— 
one from a cormorant’s stomach, the other from the thick 
coating of fat covering the abdomen—-at a meeting of the 
Royal Society of South Australia (Trans., etc., 1916, p. 590). 

Ninox rufa (No. 269).—Dr. MacGillivray found parasites under 
the skin of the head and orbit, N. Queensland (Emu. April, 
1918, p. 186). 

Podargus marmoratus (C.L. 339) (Micropodargus ocellatus mar- 
moratus).—Tapeworm in subcutaneous. tissue of the abdomen, 
N. Queensland, MacGillivray (Emu, April, 1918, p. 189). 

H. L. White, in “North Australian Birds observed by Wm. 
McLennan” (Emu, xvi., pt. iv., pp. 205-230), mentions the 
finding of the following parasites :— ; 

Turnix castanota (No. 14).—Small worms in chest, abdominal 
cavity, and eye socket. King River. 

Falco lunulatus (No. 258).—Tapeworms in a mass of yellow pus, 
and a growth containing pus and worms on the left leg. Very 
small worms in the eye socket and membrane. A short, thick, 
round worm in the abdominal cavity. A mass of long, thin, 
round worms over the kidneys and testes; the testes almost 
totally destroyed. Some of the worms over 6 ins. long. Morn- 
ington Island, July, 1915. 

Ninox boobook (No. 263).—A mass of worms in the inflamed fibrous 
membrane on the skull between the eyes; two more in the left 
eye socket and one in the abdominal cavity. ‘ 

Ninox connivens (No. 267).—Several worms under the skin of the © 
body and legs. 

Ninox rufa (No. 269).—Numérous tapeworms under the skin of 
the legs. A round worm in the tof eye. 

Trichoglossus rubritorques (No. 275).—A number of large tape- 
worms in the abdominal cavity. 

Halcyon sanctus (No. 349).—Two large and two small worms in the 
neck. One large worm in the abdominal cavity. 

Graucalus melanops (No. 457).—Small worms in the nictitating 
membrane. Large tapeworms in the intestine. 

poe hypoleucus (No. 458).—Worms under the skin of the 
thighs. 

Colluricincla brunnea (No. 568).—Small worms in the eye membrane 
and larger ones in the liver. King River. 

Colluricincla woodwardi (No. 571).—Worms under the skin. 
King River. 

Philemon sordidus (No. 685a).—A number of worms in the 
abdominal cavity. 

Oriolus flavicinctus (No. 713).—Two small worms in the abdominal 
cavity. 


Pr. II1.—ParRasites oF AUSTRALIAN BIRDS THAT HAVE COME 
UNDER THE WRITER’S NOTICE. 


1. Cestodes. 
Chaleophaps chrysochlora (No. 29).—Stradbroke Island, Q’land, 
Sept., 1919. ae 
Phaps elegans (No. 31).—Waitpinga, Encounter Bay, Jan., 1922 


105 


Podiceps poliocephalus (No. 58).—Muswellbrook, Feb. (Dr. Darnell 
Smith), cestodes in intestines, the largest in subperitoneal 
tissue(?), probably from injury. 

HKudyptula minor (No. 62).—Kncounter Bay, Feb., 1921, numerous 
cestodes in intestines, bird thin; and Jan., 1922 (2, 1: nil, 
1 with cestodes). 

Puffinus sphenurus (No. 69).—little Bay, Sydney, Dec., 1914 
(washed up dead). 

Puffinus griseus (No. 72).—Washed ashore near Manly, Oct., 1916. 

Puffinus brevicaudus (No. 74).—Flinders Island, Nov., 1912 (4 
birds, 3 nil). 

Sterna cristata (No. 107).—Encounter Bay, Jan., 1922. 

Pisobia acuminata (No. 162).—Gular, Oct., 1911 (2 birds). 

Gallinago australis (No. 166).—Mannum, S. Austr., Nov., 1913. 


- Chenopis atrata (No. 198).—In captivity, Coast Hospital, Sydney, 


April, 1916, numerous cestodes; Zool. Gardens, Sydney, Mar., 
1915. 
Anas superciliosa (No. 208).—Deniliquin, Mar., 1918 (J. Weir, per 
W. W. Froggatt); N.S. Wales (ova in tumours of intestine, 
(?) nematode or cestode, from Dr. Darnell-Smith). 
Teal.—Cobar, Dec., 1911. 


‘Hieracidea berigora (No. 259).—Flinders Island, Nov., 1912, 


cestode(?) (with nematodes in crop and stomach). 

Ninox boobook (No. 263).—Mannum, S. Austr., Nov., 19138; 
Flinders Island, Nov., 1912; Bunya Mountains, Q’land, Oct., 
1919. 

Trichoglossus swainsoni (No. 274).—Eidsvold, Q’land, July, 1913 
(Dr. T. L. Bancroft); Encounter Bay, Feb., 1921 (nil). 

Cacatua galerita (No. 291).—Sydney, in captivity, Aug., 1918. 

Barnardius barnardi (No. 315).—Willbriggie, N.S. Wales, Oct., 
1912; near Morgan, Nov., 1918 (nil); Beltana, Aug., 1921 (nil). 

Podargus marmoratus (No. 339).—Claudie River, N. Q’land, 1913 

- (Dr. MacGillivray, cestode under skin of abdomen). 

Syma. flavirostris (No. 344).—N. Q’land, 1913 (Dr. MacGillivray, 
2, cestodes in subcutaneous tissues of leg in one). 

pero macleayi (No. 347).—Stradbroke Island, Q’land, Sept., 


Pitta strepitans (No. 377).—Bunya Mountains, Q’land, Oct., 1919 
(2 with cestodes, 1 nil). 

Cheramoeca leucostenum (No. 385).—Narrabri, Feb., 1912 (2). 

Petrochelidon nigricans (No. 386).—Stradbroke Island, Q’land, 
Sept., 1919. - 

Petrochelidon ariel (No. 387).—Gular, Oct., 1911 (with trematodes) ; 
Morgan, 1913 (2 nil). 

Eopsaltria chrysorrhoa (No. 419).—Stradbroke Island, Q’land, 
Sept., 1919; Bunya Mountains, Q’land, Oct., 1919 (nil). _ 

Pachycephala melanura (No. 426).—Stradbroke Island, Q’land, 
Sept., 1919 (1 cestodes, 2 nil). 

Pachycephala rufiventris (No. 480).—Pilliga Scrub, Oct., 1918 (2, 
1 nil); Kendall, Jan., 1919 (nil); Stradbroke Island, Q’land, 
Sept., 1919; Beltana, Aug., 1912 (nil). 

Pachycephala olivacea (No. 433).—Flinders Island, Nov., 1912. 

Piezorhynchus nitidus (No. 451).—N. Q’land, 1913 (Dr. MacGil- 
livray, larval cestode (?) in subcuteaneous tissues). 

Coracina parvirostris (No. 457a).—Flinders Island, Nov., 1912 
(small bodies, (?) parasitic). © 

Hylacola pyrrhopygia (No. 474).—Encounter Bay, Jan., 1912: 
Bumberry, near Manildra, Jan., 1916 (nil). 


106 


Malurus longicaudus (No. 529). —Flinders Island, Nov., 1912 (9, 
2 with cestodes, 2 with filaria in peritoneum, 5 nil), 

Artamus leucogaster (No. 559).—Stradbroke Island, Q’land, Sept., 
1919 (1 cestodes, 1 nil). 

Artamus personatus (No. 561).—North of Renmark, Jan., 1921. 

Artamus melanops (No. 562a).—Cobar, Oct., 1911; Gunnedah, 
Sept., 1914 (mil); Beltana, Aug., 1921 (nil). 

Artamus sordidus (No. 564).—Hawkesbury River, Oct., 1912 (nil); 
Manilla, Sept., 1914 (nil); Coonabarabran, Sept., 1914; Upper 
Manilla, Sept., 1914 (nil); Bibbenluke, N.S. Wales, Mar., 
1913 (nil). 

Colluricincla harmonica (No. 566).—Hawkesbury River; June, 1912 
(nil) ; Coonabarabran, Sept., 1914 (nil) ; Encounter Bay, ‘Jan., 
SPAR 

Colluricincla selbii (No. 567).—Flinders Island, Nov., 1912. 

Corcorax melanorhamphus (No. 577).—Near 1] Morgan, Nov., 1913; 
Gunnedah, Sept., 1914 (8, echinorhynchs in 1, nil in 2); Coona- 
barabran, ‘Sept., "1914; Belaringar, April, 1915 (echinorhynchs 
and (2) cestodes) ; Tarcoon, Oct., 1914 (nil) : Dubbo, June, 1915 
(worms). 

Zosterops dorsalis (No. 599).—Sydney, June and July, 1912, and 
Nov., 1911 (all with cestodes), and Aug., 1911 (1), June, 1912 
(4), July, 1912 (11), Aug., 1912 (8), and Dec., 1918 (1) (all 
nil) : Flinders Island, Nov., 1912 (8, 2 with cestodes) ; Bunya 
Mountains, Q’land, Oct., 1919 (2 nil) ; Encounter Bay, Jan., 
1921 (nil). 

Pardalotus striatus (No. 603).—Near Morgan, Nov., 1918; 
Alawoona, S. Austr., Dec., 1918; Beltana, Aug., 1921 (nema- 
tode in peritoneum only); north of Renmark, Jan., 1921 (nil). 

Pardalotus affinis (No. 605) (Pall this species).—Flinders Island, 
Nov., 1912 (4, cestodes in 1, cestodes(?) in 1, nil in 2). 

eh albifrons (No. 630).—Overland Corner, 8. Austr., Nov., 

1913 (2, 1 nil). 

Stigmatops ocularis (No. 639).—Stradbroke Island, Q’land, Sept., 
1919 (2 with cestodes, 1 nil). 

Ptilotis fusca (No. 643). — Grafton, Ake 1912 (2 nil); Molong, Oct., 
1913 (nil); Wellington, N.S. Wales, Nov., 1914 (2 nil); Dubbo, 
July, 1914 (2 nil); French’s Forest, Sydney, June, 1915 (nil) ; 
Bumberry, near Manildra, Jan., 1916 (cestodes) ; Bumberry, 
Oct., 1916 (nil). 

Ptilotis "auricomis (No. 652).—Hawkesbury River, June, 1912; 
Molong, Oct., 1913 (nil); Grafton, April, 1912 (nil) ; Hawkes- 
bury River, April, 1913 (nil). 

Ptilotis ornata (No. 656).—Alawoona, S. Austr., Dec., 1913; 
Monarto South, May, 1921. 

Meliornis sericea (No. 669).—Stradbroke Island, Q’land, Sept., 
1919 (cestodes in 1, nil in 1). 

Myzantha flavigula (No. 674).—Tarcoon, Oct., 1914; Belaringar, 
April and May, 1915 (both nil). 

pepe rufigularis (No. 679).—Cobar, Oct., 1911. (nematodes 
only); Yaneo, Oct., 1912 (nil); Overland Corner, S. Austr., 
Nov., 19138; Narrabri, Nov. 1916 (nil). 

Entomyza cyanotis (No. 680). —Mannum, Noyv., 1913 (2 nil); Bum- 
berry, near Manildra, Jan., 1916. 

Anthus australis (No. 687). SWlinders Island, Nov., 1912 (2, 1 nil); 
West Island, Encounter Bay, Jan., 1929 (nil). 

Mirafra horsfieldi (No. 688). -_‘Wncounter Bay, Jan., 1922. 


107 


Ptilonorhynchus holosericeus (No. 718).—Bunya Mountains, Q’land, 
_ Oct., 1919 (cestodes in 1, nil in 5). 

Ailuroedus smithi (No. 720).—Mummulgum, near Casino, Dec., 
1916; Bunya Mountains, Q’land, Oct., 1919 (2 nil, 2 with 
_cestodes). 

Sericulus chrysocephalus (No. 726).—Bunya Mountains, Q’land, 
Oct., 1919 (cestode in 1, cestode in abdominal cavity (probably 
from wound) in 1, nil in 4); Mummulgum, near Casino, Dec., 
1916 (nil); Zool. Gardens, Sydney, Nov., 1919 (nil). * 

Corvus coronoides (No. 732) and C. australis (No. 734).—Yanco, 
Oct, 1912; Flinders Island, Nov., 1912 (1 nil, (?) cestode in 1); 
Walgett, Sept., 1914 (2 with cestodes); Upper Manilla, Sept., 
1914 (nil) ; Coonabarabran, Sept., 1914 (nil); Moree, Oct., 1914 
(2 nil); Tarcoon, Oct., 1914 (nil); Merah, near Moree, Oct., 
1914 (2 nil); Belaringar, June, 1915 (8, cestodes in 1); Tarcoon, 
foe 1914 (?sparganum); Bumberry, near Manildra, Jan., 

Corvus cecilae.—Stradbroke Island, Q’land, Sept., 1919 (cestodes 
on, to nil in: 2). 

Strepera graculina (No. 735).—Mount Irvine, June, 1915 (filaria 
only); Scone, May, 1917 (filaria in 1, nil in 1); Bunya Moun- 
tains, Q’land, Oct., 1919 (cestodes only). 


2. Nematodes. 


EKudyptula minor (No. 62).—Encounter Bay, Feb., 1921 (in 
stomach), and Jan., 1922 (2, no nematodes). 

Pelagodroma marina (No. 65).—Flinders Island, Nov., 1912 (6, 
nematodes in crop of 1, 5 nil). 

Pisobia acuminata (No. 162).—Flinders Island, Nov., 1912 (? nema- 
tode in intestine). 

Pelecanus conspicillatus (No. 233).—Sep., 1918 (nematodes in 
stomach). 

Astur novae-hollandiae (No. 237).—N. Queensland (Dr. MacGil- 
livray), 1913 (nematode in nictitating membrane of eye). 
Falco lunulatus (No. 258).—Flinders Island, Nov., 1912 (filaria in 

peritoneal cavity). 

Hieracidea berigora (No. 59).—Flinders Island, Nov., 1912 (nema- 
todes in stomach and oesophagus, (?)cestodes also). 

Ninox rufa (No. 269).—N. Queensland (Dr. MacGillivray), 1913 
(3, in orbit of one, orbit and under skin of forehead in 
another, 2 large flesh-coloured worms in abdominal cavity, 
and 1 small white worm in chest cavity in third). 

Pseudopsittacus maclennani.—N. Queensland (Dr. MacGillivray), 
1913 (nematodes in abdominal cavity). 

Dacelo gigas (No. 345).—Pilliga Scrub, Oct., 1918 (large nematode 
in intestine). 

Haleyon sanctus (No. 349).—Stradbroke Island, Moreton Bay 
(?nematodes in intestine and small coiled nematode in peri- 
toneal cavity, from injury to intestine). 

Pitta mackloti (No. 378).—N. Queensland (Dr. MacGillivray), 1918. 

Petroica phoenicea (No. 393).—Flinders Island, Nov., 1912 (filaria . 
in peritoneal cavity). — 

Myiagra plumbea (No. 444).—Stradbroke Island, Q’land, Sept., 
1919. 


Pomatorhinus superciliosus (No. 479).—Baradine, Oct.,_ 1918 
(nematode in intestine, also Echinorhynchus pomatostomi, sub- 
cutaneously). 


108 


Oreocincla macrorhyncha (No. 4884).—Mount Arthur, near Laun- 
ceston, Tas., Nov., 1912 (Pnematode in intestine). 

Malurus longicaudus (No. 529).—Flinders Island, Nov., 1912 (9, 
filaria in peritoneum of 2 birds, cestodes in 2, nil in 5). 

Pardalotus striatus (No. 603).—See under .Cestodes. 

Ptilotis leilavalensis (No. 6614).—Beltana, Aug., 1921 (nematode 
attached to outer wall of oesophagus). 

Myzantha flavigula (No. 674).—North of Renmark, Jan., 1921 
(nematodes in pleuro-peritoneal cavity, yellow, as was the 
fat and skin of the abdomen. 

Acanthogenys rufigularis (No. 679).—Cobar, Oct., 1911 (nematode 
only); Yanco, Oct., 1912 (nil); Overland Corner, S. Austr., 
Nov., 1912 (cestode only); Narrabri, Nov., 1916 (nil). 

Strepera graculina (No. 735).—Mount Irvine, June, 1915 (filaria 
in peritoneal cavity); Scone, May, 1917 (filaria in pleuro- 

_ peritoneal cavity in 1, nil in 1); Bunya Mountains, Q’land, 
Oct., 1919 (cestodes only). 

Domestic pigeons, chiefly squabs about 28 days old.—Numerous 
small nematodes in intestine, almost blocking it, Sydney, May, 
1919; Ascaridea. columbae (Gmelin) (Heterakis maculosa, 
Schn.), identified by Miss Irwin Smith. 


3- Acanthocephala. 


Baza subcristata (No. 254).—_Mummulgum, N.S. Wales, Dec., 1916, 
Centrorhynchus asturinus, Jnstn. 

Seisura inquieta (No. 448).—Canowindra, 1915 (echinorhynch near 
rectum). 

Cinclosoma cinnamoneum (No. 468).—(?) Locality, larval Echinor- 
hynchus pomatostomi, C. and J., in subcutaneous tissue of 
neck (Dr. MacGillivray). 

iri crepitans (No. 476).—Bunya Mountains, Q’land, Oct., 


Pomatorhinus temporalis (No. 478).—Canowindra, 1915 (3, larval 
E. pomatostomi). 

Pomatorhinus superciliosus (No. 479).—Hallett Cove, S. Ausir., 
May, 1910, larval E. pomatostomi subcutaneously ; Baradine, 
Oct., 1918, larval E. pomatostomi subcutaneously (also nema- 
todes in intestine). . 

Oreocincla lunulata (No. 488).—Bunya Mountains, Q’land, Oct., 
1919 (echinorhynchs in 5); Kuitpo, S. Austr., May, 1921, 
larval E. pomatostomi subcutaneously. 

Corcorax melanorhamphus (No. 577).—Near Morgan, Nov., 1913 
(cestode only); Gunnedah, Sept., 1914 (8, echinorhynch in 1); 
Coonabarabran, Sept., 1914 (cestode only); Belaringar, April, 
1915 (echinorhynchs and cestodes ?); Tarcoon, Oct., 1914 (nil); 
Dubbo, June, 1915 (worms). 

Aphelocephala leucopsis (No. 578).—Hallett Cove, S. Austr., May, 
1910, larval E. pomatostomi subcutaneously ; Gular, Oct., 1911 
(2 nil); Narrabri, Feb., 1912 (nil); Overland Corner, S. Austr., 
Dec., 1913 (nil); Mount Lofty Ranges, Nov., 1912 (nil); north 
of Renmark, Jan., 1921 (nil). : 

Climacteris scandens (No. 592) (C. picumnus).—Near Morgan, 
Nov., 1913 (2, larval E. pomatostomi and (?)worm in intes- 
tine in 1, nil in 1). 


109 


4. Trematodes. | 


Petrochelidon ariel (No. 387).—Gular, Oct., 1911, type of Plagiorcis 
clelandi, S. J. Johnston (with cestodes in intestines); Morgan, 
Nov., 1913 (2, both nil). 


5. Species of Birds Examined in which Entozoa 
(excluding Haematozoa) have not been 
detected by the Writer. 


Leipoa ocellata (No. 6).—Zool. Gardens, Sydney (2 birds). 

Coturnix pectoralis (No. 8).—Encounter Bay, Jan., 1922 (2). 

Turnix varia (No. 13).—Flinders Island, Nov., 1922. 

Turnix velox (No. 16).—Near Broken Hill, April, 1917. 

oes humeralis (No. 26).—Stradbroke Island, Q’land, Sept., 

Geopelia tranquilla (No. 27).—Coonamble, Aug., 1912; Mannum, 
S. Austr., Nov., 1918 (2). . 

Bee ee cortors (No. 30).—Overland Corner, S. Austr., Dec., 
9 


Ocyphaps lophotes (No. 39).—Parachilna, Aug., 1921. 

Leucosarcia picata (No. 40).—Bunya Mountains, Q’land, Oct., 1919. 

Haematopus fuliginosus (No. 126).—Flinders Island, Nov., 1912. 

Lobivanellus lobatus (No. 128).—Upper Manilla, Sept., 1914. 

Spoonbill (white).—Taronga Zool. Park, June, 1919. 

Astur novae-hollandiae (No. 237).—Taronga Zool. Park, April, 1919. 

Astur approximans (No. 228).—N.S. Wales, April, 1912. 

Uroaétus audax (No. 243).—Nevertire, Aug., 1919. 

Haliastur sphenurus (No. 248).—Coonamble, Aug., 1912; Tarcoon, 
Oct., 1914. 

Calyptorhynchus baudini (No. 284).—Taronga Zool. Park (from 
W. Austr.), Aug., 1919. 

Calyptorhynchus leachi (No. 289).—Narrabri, Nov., 1916 (2 birds) ; 
Dorrigo, Jan., 1918. 

Cacatua gymnopis (No. 293).—Beltana, Aug., 1921. 

Cacatua roseicapilla (No. 295).—Belaringar, April, 1915. 

Aprosmictus scapulatus (No. 303).—Bunya Mountains, Q’land, 
Oct., 1919 (8). 

Platycercus pennanti (No. 304).—Wagga, July, 1914 (2); Mount 
Irvine, June, 1915; Bunya Mountains, Q’land, Oct., 1919 (7). 

Platycercus flaveolus (No. 306).—Morgan, S. Austr., Nov., 1913 (3). 

Platycercus flaviventris (No. 307).—Flinders Island, Nov., 19138 (4), 

Platycercus eximius (No. 311).—Belaringar, May, 1915; Dubbo, 
July, 1915 (2); Kendall, Aug., 1918. 

Psephotus haematogaster (No. 319).—Belaringar, May, 1915. 7 

Psephotus multicolor (No. 323).—North of Renmark, Jan., 1921 (2). 

Psephotus haematonotus (No. 324).—Cowra, Sept., 1911; Coon- 
amble, Aug., 1912; Mannum, S. Austr., Nov., 1913 (2); Goolwa, 
Nov., 1921. 

Euphema elegans (No. 327).—Encounter Bay, Jan., 1921, and Jan., 
1922 


Euphema pulchella (No. 330).—Narrabri, June, 1919. 

Lathamus discolor (No. 332).—Flinders Island, Nov., 1912. 

Eurystomus pacificus (No. 341).—Scone, Oct., 1917. ‘3 

Halcyon pyrrhopygius (No. 348).—Near Morgan, S. Austr., Nov., 
1913. 5 . 


Merops ornatus (No. 352).—Coonabarabran, Sept., 1914. 
- Cuculus pallidus (No. 361).—N.S. Wales, Nov., 1911. 


110 


Cacomantis flabelliformis (No. 362).—Flinders Island, Nov. 1913 (2). 

Chalcococcyx basalis (No. 366). ee anor Island, Nov., 1913; Over- 
land Corner, S. Austr., Dec. 

Eudynamis cyanocephala (No. shi) ae a near Casino, 
Dec., 1916 (2 birds). 

Microeca fascinans (No. 388).—Sydney, Nov., 1911; Morgan, 
S. Austr., Nov., 

Petroica legeii (No. 392). —Flinders Island, Nov., 1911. 

Petroica phoenicea (No. 393).—Flinders Island, 'Nov., Loi 

Petroica goodenovii (No. 394).—Beltana, Aug.., 1922. 

Erythrodryas rosea (No. 396). —Hawkesbury River, June, 1912. 

Melanodryas bicolor (No. 397).—Pilliga Scrub, Oct.., 1918 ; 
Encounter Bay, Jan., 1921. 

Amaurodryas vittata (No. 398).—Flinders Island, Nov., 1911 (2). 

Orthonyx ‘spinicaudis (No. 464).—Dorrigo, Jan., "1918 (2 birds). 

Smicrornis brevirostris (No. 400). —Morgan, Nov., 1913; Scone, 
May, 1917; Dubbo, Aug., 1917; Pilliga Scrub, Oct., 1918 ; 
north of Renmark, Jan., 1921. 

Gerygone albogularis (No. 402). —Molong, Oct., 1913. 

Gerygone fusca (No. 405).—Lisarow, May, 1915. 

Kopsaltria australis (No. 418). —Molong, Oct., 1913. 

Falcunculus frontatus (No. 422). —Mount Irvine, June, 1915. 

Oreoica cristata (No. 425).—North of Renmark, di an., 1921. 

Pachycephala gutturalis (No. 428). orca Mountains, Q’land, 
Oct., 1919 (2); Encounter Bay, Jan. 

Pachycephala elaucura (No. 429). Flinders and Nov., 1912 (8). 

Rhipidura diemenensis (No. 436a).—Flinders Island, Nov., 1912 (8). 

Rhipidura rufifrons (No. 489).—Mummulgum, near Casino, Dec., 
1916; Bunya Mountains, Q’land, Oct., 1919. 

Rhipidura motacilloides (No. 442). __Sydney, Nov., 1911. 

ee carinata (No. 455).—Bunya Mountains, Q’land, Oct., 


Graucalus melanops (No. 457).—Tarcoon, Oct., 1914; Upper 
Manilla, Sept., 1914; Beltana, Aug., 1921. 
Graucalus parvirostris (No. 4574).—Flinders Island, Nov., 1912 (4). 


~ Graucalus mentalis (No. 459).—Coonabarabran, Sept., 1914 


Campephaga humeralis (No. 462).—Hawesbury River, Oct., 1912; 
Baan Baa, Jan., 1917 (young bird). 

Campephaga leucomela (No. 463).—Stradbroke Island, Q’land, 
Sep., 1919. 

Cinclosoma punctatum (No. 466).—Encounter Bay, Jan., 1922. 

pare eager se (No. 467).—Alawoona, S. Austr., Dec., 

rth brunneopygius (No. 472).—Alawoona, S. Austr., Dec., 

Hylacola cauta (No. 475).—Monarto South, S. Austr., May, 1921. 

Cincloramphus-cruralis (No. 484).—Near Broken Hill, April, 1917. 

Cincloramphus rufescens (No. 485).—Pilliga Scrub, Oct., 1918 (2). 

Ephthianura albifrons (No. 489).—Flinders Island, Nov., 1913; 
Encounter Bay, Jan., 1921 (2). 

Ephthianura tricolor (No. 490).—Molong, Oct., 1913 (8); Para- 
chilna, S. Austr., Aug., 1921. 

Ephthianura aurifrons (No. 491).—Broken Hill, April, 1917; 
Parachilna, S. Austr., Aug., 1921. 

Origma rubricata (No. 500).—Sydney, April, 1912. 

Chthonicola sagittata (No. 501).—The Oaks, N.S. Wales, June, 
1914; Baan Baa, near, Boggabri, Jan., 1917. 


vit 


Acanthiza nana (No. 503).—Hawkesbury River, May, 1915; Dubbo, 
Mar., 1915; Pilliga Serub, Oct., 1918; Narrabri, June, 1919; 
Bunya Mountains, Oct., 1919. 

Acanthiza reguloides (No. 507). —Bibbenluke, Mar., 1913; Pilliga 
Serub, Oct., 1918; Bunya Mountains, Q’land, Oct., 1919. 

Acanthiza chrysorrhoa (No. 508). —Scone, May, 1917. 

Acanthiza uropygialis (No. 509). Vanco, Oct., 1912: Mannum, 
Nov., 1913; Overland Corner, S. Austr., Dec., 1918; Dubbo, 
July, 1915 (2): Baan Baa, Jan., 1917; Beltana, Aug., 1921 ; 
north of Renmark, Jan., 1921. 

Acanthiza lineata (Not 511).—Sydney, Nov., 1912; Bell, June, 
1915; Bunya Mountains, Oct., 1919; Encounter Bay, Jan., 
1921, and Jan., 1922. 

Acanthiza pusilla (No. 512).—Kurrajong, Aug., 1912; Bibbenluke, 
N.S. Wales, Mar., 1913; Bunya Mountains, Q’land, Oct.. 
1919 (2). 

Acanthiza diemenensis (No. 512a).—Flinders Island, Nov., 1913 (2). 

Acanthiza pyrrhopygia (No. 516).—Monarto South, S. Austr., 
July, 1914; Encounter Bay, Feb., 1921. 

Acanthiza albiventris (No. 5164).—Pilliga Scrub, Oct., 1918. 

Pyrrholaemus brunneus (No. 517).—Renmark, Jan., 1921. 

Sericornis citreigularis (No. 518).—Mount Irvine, June, 1915; 
Bunya Mountains, Q’land, Oct., 1919 (2). 

Sericornis frontalis (No. 519). = Tisarow, May, 1915 (2); Mount 
Irvine, June, 1915; Canobolas, Oct., 1916 (2); Bunya Moun- 
tains, 'Q’land, Oct., 1919. 

Roos magnirostris (No. 521).—Bunya: Mountains, Q’land, 

ct., : 

Sericornis humilis (No. 524).—Flinders Island, Nov., 1912 (5). 

Malurus cyaneus (No. 5380).—Sydney, Nov., 1911; Kuitpo, S. Austr., 
May, 1921. 

Malurus cyanochlamys (No. 530a).—Bunya Mountains, Q’land, 
Oct., 1919 (5). 

aarus ‘melanonebus (No. 5382).—Overland Corner, S. Austr., Dee., 

13 

Malurus cyanotus (No. 535).—Beltana, Aug., 1921. 

Malurus assimilis (No. 538).—Alawoona, S. Austr., Dec., 1913 (2); 
Beltana, Aug., 1921. 

Malurus melanocephalus (No. 542).—Mummulgum, near Casino, 


Dec., 1916. 

Artamus superciliosus (No. 560).—Cowra, Sept., 1911; Sydney, 
Oct., 1919. 

Su rufigaster (No. 573).—Stradbroke Island, Sept., 
1914 (4 


Grallina picata (No.. 575).—Cowra, Sept., 1911; Pennant Hills, 
Sydney, Dec., 1916 (D. Steel). 

Struthidea einenes (No. 576).—Gunnedah, Sept., 1914 (3); Coona- 
barabran, Sept., 1914; Belaringar, April, 1915. 

Neositta chrysoptera (No. 583).—Hawkesbury River, June, 1912. 

Neositta pileata (No. 586).—Encounter Bay, Feb., 1921. 

Climacteris picumna (No. 592) (scandens). ie arrabri, Feb., 1912; 
Molong, Oct., 1913; Baradine, Oct., 

peers lencophaea (No. 593). Lites Ren ee ts Q’ land, Oct., 


Pardalotus punctatus (No. 606).—Flinders Island, Nov., 1912. 
Pardalotus xanthopygius (No. 607).—Mannum, Nov., 1913. 


112 


Pardalotus melanocephalus (No. 609).—Stradbroke Island, Q’land, 
Sept., 1919. 

Melithreptus lunulatus (No. 613). __Sydney, Nov., 1911 (2), and 
May, 1912; Hawkesbury River, June, 1912 (2) ; Stradbroke 
Island, Q’land, Sept., 1919; Kuitpo, S. Austr. , May, 1921. 

Melithreptus brevirostris (No. 619). Sydney, April, 1912; 
Hawkesbury River, June, 1912; Mannum, S. Austr., Nov., 
1913 (3); Scone, May, 1917: Bumberry, Oct., 1916 ; Encounter 
Bay, Jan., 1921. 

A it melanocephalus (No. 620).—Flinders Island, Nov., 


Myzomela sanguineolenta (No. 622).—Kendall, Jan., 1919. 

Myzomela nigra (No. 624).—Molong, Oct., 1913. 

Glycyphila fulvifrons (No. 629).—Flinders “Island, Nov., 1912 (4); 
French’s Forest, Sydney, June, 1915. 

Meliphaga phrygia (No. 638). - Bumberry, Sept., 1916. 
Ptilotis chrysotis (No. 644).—Bunya Mountains, Q’land, Nov., 1919. 
Ptilotis sonora (No. 646).—Mannum, S. Austr., Nov., 1913; Para- 
chilna, S. Austr., Aug., 1921; Encounter Bay, Jan., 1922. 
Ptilotis chrysops (No. 648). __Hawkesbury River, June, 1912; 
Kurrajong, Aug., 1912; Hawkesbury River, Nov., 1914, and 
May, 1915. 

Ptilotis flavigula (No. 649).—Flinders Island, Nov., 1912 (8). 

Ptilotis leucotis (No. 651).—Dubbo, Aug., 1917. . 

Ptilotis ornata (No. 656). Cerca Nov., 1913 (2); Monarto 
South, S. Austr., July, 1914. 

Ptilotis plumula (No. 658).—North of Renmark, Jan., 1921. 

Ptilotis penicillata (No. 661).—Narrabri, Feb., 1912; near Morgan, 
S. Austr., Nov., 1913 (2); Overland Corner, S. Austr., Dec., 
I9IS: Pilliga Serub, Oct.., 1918. 

Lichmera australasiana (No. 667). —Flinders Island, Nov., 1912. 

Myzantha garrula (No. 672).—Gunnedah, Sept., 1914 (4); Upper 
Manilla, Sept., 1914; Hawkesbury River, May, 1915; Bela- 
ringar, April (2) and. May, 1915; Scone, May, 1917. 

Anthochaera carunculata (No. 675).__Hawkesbury River, July, 
1912 (8); Sept., 1912 (4); Scone, May, 1917. . 

a ae corniculatus (No. 684).—Hawkesbury River, May, 


eee cr onutanys (No. 685).—Cowra, Sept., 1911; Dubbo, 
. 
Panera Phe bellus (No. 693).—Flinders Island, Nov., 1912. 
Stizoptera bichenovii (No. 697).—Narrabri, June; 1919 (3). 
Aegintha temporalis (No. 703).—Gosford, May, 1915 (4); Encounter 
Bay, Jan., 1922. 
ance note (No. 728).—Bunya Mountains, Q’land, Oct., 
Strepera arguta (No. 736).—Flinders Island, Nov., 1912 (2). 
Cracticus destructor (No. 745).—Tarcoon, Oct., 1914. 
Gymnorhina tibicen (No. 747).—Upper Manilla, Sept., 1914; Tar- 
coon, Dec., 1914. 
eins patzaeo (introduced Dove).—Sydney, Nov., 1911, and Mar., 


Sturnus ‘vulgaris (Starling).—Gunnedah, Sept., 1914 (5, young); 
Wagga, Aug., 1914 (2). 


Passer domesticus (Sparrow).—Sydney, Nov., 1911, and June, 1917. 


1]3 


6. Siphonaptera (Fleas). 


Eudyptula minor (No. 62).—Bird Island, Rockingham, W. Austr., 
“Hh 1906 (Parapsyllus australiacus, Rothsch., Nov. Zool., 
XV1., 1909, p. 62, wm cop.); Flinders ‘Island, Nov., 1912 (P. 
australiacus, determined by ic, Rothschild) : Encounter Bay, 
Feb., 1921 (no fleas, mallophaga), and Jan., 1922 (no fleas, 
mallophaga). 

Puffinus brevicaudus (No. 74).—Flinders Island, Nov., 1912 (P. 
australiacus, Rothsch.; doubtful as to whether a str ay). 


7. Diptera. 


Pokehelidan ariel (No. 387).—Near Morgan, S. Austr., Nov., 1913 
Ornithomyia australasiae Leach(? ), identified at British 
Museum). 


8. Waitephasn: 


Coturnix pectoralis (No. 8).—Encounter Bay, Jan., 1911 (1 nil, 
1 mallophaga on wings). 

Turnix velox (No. 16).—Near Broken Hill, April, 1917. 

Phaps elegans (No. 31).—Waitpinga, Encounter Bay, Jan., 1922. 

Eudyptula minor (No. 62).—See under Siphonaptera. 

Pelagodroma marina (No. 65).—Flinders Island, Nov., 1912 (1 
mallophaga, 1 nil). 

Puffinus sphenurus (No. 69).—little Bay, Sydney, Dec., 1914 
(washed ashore). 

Puffinus brevicaudus (No. 74).—Flinders Island, Nov., 1912 (2). 

Prion banksi (No. 89).—Cronulla, Aug., 1911 (washed up). 

Sterna cristata (No. 107).—Encounter Bay, Jan., 1922. 

Haematopus fuliginosus (No. 126).—Flinders Island, Nov., 1912 
(1 mallophaga, 1 nil). 

Lobivanellus lobatus (No. 128).—Upper Manilla, Sept., 1914 
(mallophaga and mites). 

Himantopus leucocephalus (No. 142).—(?) Locality (Dr. D’Ombrain). 

Pisobia acuminata, (No. 162).—Gular, Oct., 1911; Flinders Island, 
Nov., 1912 (nil); Cape York or south-west of Queensland, 
Dec.. 1912 (Dr. MacGillivray). 

Rhynchaea australis (No. 167). Cape York or south-west of Queens- 
land, Dec., 1912 (Dr. MacGillivray). 

Chenopis atrata (No. 198).—In captivity, Coast Hospital, Sydney, 
April, 1916; Zool. Gardens, Sydney. 

Cereopsis novae-hollandiae (No. 202).—Cape Barren Island, Bass 
Straits, Nov., 1912. 

Astur approximans (No. 238).—N.S. Wales, April, 1912. 

Uroaétus audax (No. 243).—Nevertire, Aug., 1919. 

Haliastur sphenurus (No. 248).—Coonamble, Aug., 1912; Tarcoon, 
Oct., 1914. 

Kestrel—From Dr. D’Ombrain. 

Hieracidea berigora (No. 259).—Flinders Island, Nov., 1912. 

Hieracidea occidentalis (No. 260).—Narrabri, Jan-, 1918. 

Trichoglossus swainsoni (No. 274).—Encounter Bay, Jan., 1921. 

Calyptorhynchus leachi (No. 289).—Narrabri, Nov., 1916 (2); 
Dorrigo, Jan., 1918 (nil). 

Eclectus macgillivrayi.North Queensland (Dr. MacGillivray). 

Platycercus flaviventris (No. 307).—Flinders Island, Nov., 1912. 

Platycercus eximius (No. 311).—Belaringar, May, 1915; Dubbo, 
July, 1915 (2 nil); Kendall, Aug., 1918 (nil). 


114 


Psephotus haematonotus (No. 324).—Cowra, Sept., 1911 (nil) ; Coon- 
amble, Aug., 1912. 

Lathamus ’ discolor (No. 332).—Flinders Island, Nov., 1912. 

Kurystomus pacificus (No. 341).—Scone, Oct., 1917. 

Pen: macleayi (No. 347). — Stradbroke Island, Q’ land, Sept. 

Merops ornatus (No. 352).—Coonabarabran, Sept., 1914. 

EKudynamis cyanocephala (No. 371). —Mummulgum, near Casino, 
Dec., 1916 (2). 

Erythrodryas rosea (No. 396).—Hawkesbury River, June, 1912. 

Monarcha carinata (No. 455).—Ourimbah, Nov., 1911. 

Graucalus melanops (No. 457).—Upper Manilla, Sept., 1914; Tar- 
coon, Oct., 1914 (2 nil); Beltana, Aug., 1921 (nil). 

Graucalus parvirostris (No. 4574). — Flinders Island, Nov., 1912 (2). 

Graucalus mentalis (No. 459).—Coonabarabran, Sept.. 1914. 

Oreocincla lunulata (No. 488).—Bunya Mountains, Q’land, Oct., 
1919; Kuitpo, S. Austr., May, 1921 (nil). 

Chthonicola sagittata (No. 501). —The Oaks, June, 1914. 

Acanthiza nana (No. 503).—The Oaks, June, 1914 (nil); Bunya 
Mountains, Q’land, Oct., 1919. 

Acanthiza reguloides (No. 507).—The Oaks, June, 1914 (2). 

Acanthiza chrysorrhoa (No. 508).—The Oaks, June, 1914; Dubbo, 
ane 1915 (nil); Scone, May, 1917 (nil) : Beltana, Aug. cf 1921 
nil 

Acanthiza uropygialis (No. 509).—Dubbo, ee 1911 (2), and July, 
1915 (2 nil); Beltana, Aug., 1921 (nil). 

Acanthiza lineata (No. 511). —Mount Irvine, June, 1915 (nil); 
Uralla, June, 1915 (nil); Encounter Bay, J an., 1921 (nil), and 
Jan., i922 (mallophaga). 

Artamus sordidus (No. 564).—Coonabarabran, Sept., 1914 (mallo- 
phaga, no mites); Hawkesbury River, Oct., 1912 (nil); Upper 
Manilla, Sept., 1914 (mites, no mallophaga). 

Colluricincla ’ harmonica (No. 566).—Hawkesbury River, June, 
1912 (mallophaga, no mites); Coonabarabran, Sept., 1914 (no 
mallophaga, mites) ; Encounter Bay, Jan., 1922 (mallophaga 
and mites). 

Struthidea cinerea (No. 576).—Coonabarabran, 1914 (mallophaga 
and mites); Belaringar, April, 1915 (nil); Gunnedah, 1914 (1 
with mallophaga and mites, 1 with mites, 1 nil). 

Corcorax melanorhamphus (No. 577). —Coonabar abran, 1914 (mallo- 
phagia, no mites); Belaringar, April, 1915 (numerous nits, oue 
mallophaga) ; Tarcoon, Oct., 1914 (mallophaga) ; Gunnedah, 
1914 (nil); Dubbo, July, 1915 (mallophaga, (?)two species, no 
mites). 

Melithreptus lunulatus (No. 613).—Sydney, Nov., 1911; Hawkes- 
bury River, June, 1912 (no mallophaga, mites) ; Abbotsford, 
Sydney, Nov., 1911 (Melithreptus, probably M. lunulatus) ; 
Sydney, April, "1919 (no mallophaga, mites); Kuitpo, S. Austr., 
May, 1921 (nil). 

Melithreptus brevirostris (No. 619).—N.S. Wales; Scone, May, 
1917; Encounter Bay, Jan., 1921 (nil). 

Myzomela sanguineolenta (No. 622). —Sydney, Oct., 1919 (Dr. 
D’Ombrain); Kendall, Jian., 1919 (nil). 

Ptilotis auricomis (No. 652). —Hawkesbury River, June, 1912. 

Myzantha garrula (No. 672).—Belaringar, April, 1915 (2, 1 with 
mallophaga), and May, 1915 (nil); Gunnedah, 1914 (4 with 
mites and no mallophaga); Upper Manilla, Sept., 1914 (mal- 
lophaga and no mites); ‘Cobar, Nov., 1911 (perhaps M. 
flavigula); Scone, May, 1917. 


_115 


Myzantha flavigula (No. 674).—Belaringar, April, 1915 (mai- 
lophaga) and May, 1915 (nil); Tareoon, Oct., 1915 (nil). 
Anthochaera .carunculata (No. 675).—Hawkesbury River, July, 
~ 1912 (4, mallophaga and mites in 1, mallophaga only in 3); 
Scone, May, 1917 (nil). : 

Philemon citreogularis (No. 685).—Cowra, Sept., 1911; Gular, 
Oct., 1911. 

Ptilonorhynchus holosericeus (No. 718).—Bunya Mountains, 
Q’land, Oct., 1919. 

re smithi (No. 720).—Mummulgum, near Casino, Dec., 


Sericulus chrysocephalus (No. 726).—Bunya Mountains, Q’land, 
Oct., 1919; Zool. Gardens, Sydney, Nov., 1919. 

Corvus coronoides (No. 732).—Cobar, Nov., 1911; Belaringar, 
June, 1915 (3, all with mallophaga and 1 with mites also); 
Coonabarabran, Sept., 1914 (mallophaga and mites); Upper 
Manilla, Sept., 1914 (mallophaga and mites); Tarcoon, Oct., 
1914 (2, 1 mallophaga only, 1 nil). 

Corvus cecilae (No. 733).—(?) Locality (Dr. MacGillivray). 

Strepera graculina (No. 735).—Scone, May, 1917 (2, mallophaga 
ores! ‘on. 1): 

Strepera arguta (No. 736).—Flinders Island, Nov., 1912. 

Gymnorhina: tibicen, (No. 747).—Cobar, Nov.. 1911 (either G. 
tibicen or G. leuconota); Upper Manilla, Sept., 1914 (mallo- 
phaga and mites); Tarcoon, Oct., 1914 (nil). 

Sturnus vulgaris (English Starling).—Wagga, Aug., 1914. 


9g. Ticks. 


_ Eudyptula minor (No. 62).—Rockingham, W. Austr., Nov., 1906 


Ornithodorus taljae (Guérin-Méneville) ? (larvae) and Ixodes 
percavatus, Neum., identified by Nuttall and Warburton; 
Flinders Island, Bass Straits, Nov., 1912; Encounter Bay, 
Feb., 1921 (mallophaga only). 

Pitta strepitans (No. 377).—Bunya Mountains, Q’land, Oct., 1919, 
Ixodes holocyclus, Newm., round head. 

Petrochelidon ariel (No. 387).—Bumberry, N.S. Wales, Oct., 1916, 
Argas lagenoplastis, Frogg., in nests. 

Sericornis citreigularis (No. 518).—Bunya Mountains, Q’land, Oct., 
Ixodes holocyclus, Newm., round head. . 


10. Mites. 


Lobivanellus lobatus (No. 128).—See under Mallophaga. 

Pachycephala gutturalis (No. 428).—Uralla, June, 1915 (nil); 
Encounter Bay, Jan., 1922 (mites on wings). 

Orthonyx spinicaudus (No. 464).—Dorrigo, Jan., 1918 (2). 

Haga punctatum (No. 466).—Encounter Bay, Jan., 1922 (on 
wings). 

Pomatorhinus temporalis (No. 478).—Canowindra, 1915 (3, red 
mites on 1). 

Origma rubricata (No. 500).—Sydney, April, 1912. 

Malurus longicaudus (No. 529).—Flinders Island, Nov., 1912. 

Malurus cyanotus (No. 535).—Beltana, Aug., 1921 (under wings). 

Artamus sordidus (No. 564).—See under Mallophaga. 

Colluricincla harmonica (No. 566).—See under Mallophaga. 

Neositta chrysoptera (No. 583).—Hawkesbury River, June, 1912. 

Climacteris scandens (picumna) (No. 592).—Narrabri, Feb., 1912. 


. 


116 


Zosterops dorsalis (No. 599).—Sydney, Aug., 1911 (nil), June, 1912 
| (4, 2 nil), July, 1912 (12, 11 nil), Aug., 1912 (8 nil), and Dec., 
1918 (1 nil). 
| Melithreptus lunulatus (No. 613).—See under Mallophaga. 
Myzantha garrula (No. 672).—See under Mallophaga. 
| | Anthochaera carunculata (No. 675).—See under Mallophaga. 
Philemon citreogularis (No. 685).—Dubbo, Aug., 1917. 
Zonaeginthus bellus (No. 693).—Flinders Island, Nov., 1912. «- 
Aegintha temporalis (No. 703).—Gosford, May, 1915 (4 with mites) ; 
Encounter Bay, Jan., 1922, nil. 
Corvus coronoides (No. 732). —See under Mallophaga. 
Cracticus destructor (No. 745).—Tarcoon, Oct., 1914. 
Gymnorhina tibicen (No. 747).—See under Mallophaga. 
Sturnus vulgaris (English Starling).—Gunnedah, 1914 (2, young, 
1 with mites). 


. No Ectozoa Detected. 
Geopelia pou Te ya 27).—Coonamble, Aug., 1912. 
Ochyphaps lophotes (No. 39). —Parachilna, Aug., 1921. 
} Ninox boobook (No. 263).—Flinders Island, Nov., 1912; Mannum, 


ee 


S. Austr., Nov., 1913. 
Cacatua gymnopis (No. 293).—Beltana, Aug., 1921. 
Cacatua roseicapilla (No. 295). —Belaringar, "April, 1915. 
Platycercus pennanti (No. 304).—Mount Irvine, June, 1915. 


| | Psephotus haematogaster (No. 319). -—Belaringar, May, 1915. 
Kuphema elegans (No. 327).—Encounter Bay, January, 1922. 
i Cuculus pallidus (No. 361).—Upper Manilla, Sept.. 1914. 
| Cacomantis flabelliformis (No. 362).—N.S. Wales, Nov., 1911. 
| Microeca fascinans (No. 388).—Sydney, Nov., 1911. 
‘ Petroica phoenicea (No. 398). —Flinders Island, Nov., 1912. 
| Petroica goodenovii (No. 394).—Beltana, Aug., 1 21. 
' Melanodryas bicolor (No. 397). —Encounter Bay, Jan., 1921. 
| 


ee brevirostris (No. 400).—Scone, May, 1917; Dubbo, Aug., 
191 


Gerygone fusca (No. 405).—Lisarow, May, 1915 
| | ner Hoperecr a Og (No. 430). —Kendall, Jan. , 1919; Beltana, 
ug., 
| | Myiagra plumbea (No. 444).—Hawkesbury River, Oct., 1912. 
i . Campephaga humeralis (No. 462).—Hawkesbury River, Oct., 1912. 
i Hylacola pyrrhopygia (No. 474).—Encounter Bay, Jan., 1921. 
| Hylacola cauta (No. 475).—Monarto South, May, 1921. 
. Cincloramphus cruralis (No. 484).—Near Broken Hill, April, 1917. 
i Ephthianura albifrons (No. 489).—Encounter Bay, Je an., 1921 (2). 
Z| Ephthianura tricolor (No. 490).—Parachilna, Aug., 1921. 
‘a Ephthianura aurifrons (No. 491).—Broken Hill, April, 1917; Para- 
+ | chilna, Aug., 1921. 
4 Sericornis frontalis (No. 519).—Lisarow, May, 1915 (2). 
hi | Mote renee (No. 530).—Sydney, Nov., 1911; ; Kuitpo, S. Austr., 
i} ay, 
i | Malurus assimilis (No. 538).—Beltana, Aug., 1921. 

Artamus superciliosus (No. 560).—C owra, Sept., 1914, 
Sar heh melanops (No. 5624). Gunnedah, 1914; Beltana, Aug., 
D | Grallina picata (No. 575).—Cowra, Sept., 1911; Pennant Hills, 
Hi Sydney, Dec., 1916 (D. Steel). 
| Aphelocephala leucopsis (No.. 578).—Gular, Oct., 1911 (2); Yanco, 
) Oct., 1912; Mount Lofty Ranges, Nov. 0. 
q . Weitere striatus (No. 603).—North of Renmark, Jan., 1921. 


ies 4a7 


Pardalotus affinis (No. 605).—Flinders Island, Nov., 1912. 

pee obus xanthopygius (No. 607).—Mannum, S. Austr., Nov., 

Glyciphila fulvifrons (No. 629).—French’s Forest, near Sydney, 
June, 1915. 

Ptilotis fusca (No. 643).—French’s Forest, June, 1915; Dubbo, 
July, 1915. 

Ptilotis sonora (No. 646).—Parachilna, Aug., 1921; Encounter 
Bay, Jan., 1922. 

Ptilotis chrysops (No. 648).—Hawkesbury River, June, 1912, and 
Nov., 1914. 

Ptilotis leucotis (No. 651).—Dubbo, Aug., 1917. 

Ptilotis ornata (No. 656).—Monarto South, May, 1921. 

_ Lichmera australasiana (No. 667).—Flinders Island, Nov., 1912. 

Ptilotis leilavalensis (No. 6614).—Beltana, Aug., 1921. 

Acanthogenys rufigularis (No. 679).—Narrabri, Nov., 1916. 

Be a, pusbralis (No. 687).—West Island, Encounter Bay, Jan., 

Mirafra horsfieldi (No. 688).—Encounter Bay, Jan., 1922. 

Turtur ferrago (introduced Dove).—Sydney, Mar., 1917. 

Passer domesticus (Sparrow).—Sydney, June, 1917. 


12. Haematozoa. 
(a) HALTERIDIA IN THE RED CORPUSCLES. 


Eudynamis cyanocephala (No. 371).—Mummulgum, near Casino, 
Dec., 1916 (halteridia in 2 with gametes in both). 

Melithreptus brevirostris (No. 619).—Encounter Bay, Feb., 1921; 
Monarto South, Oct., 1920 (nil). 

Ptilotis leilavalensis (No. 6614).—Beltana, Aug., 1921. 

Acanthogenys rufigularis (No. 679).—Narrabri, Nov., 1916 (one 
seen, occupying both ends of the red cell and one side); 
Monarto South, Oct., 1920 (nil). 

~Tropidorhynchus corniculatus (No. 684).—Milson Island, Hawkes- 

bury River, May, 1915 (with Leucocytozoon). 


(b) TRYPANOSOMES IN THE BLOOD. 
Pachycephala melaneura (No. 426).—Stradbroke Island, Q’land, 
Sept., 1919. 
Entomyza cyanotis (No. 680).—Bumberry, Jan., 1916 (one degener- 
ated trypanosome seen, with Leucocytozoon). 


(c) LEUCOCYTOZOA IN THE BLOOD. 


Entomyza cyanotis (No. 680).—Bumberry, Jan., 1916 (with 
trypanosomes). 

Tropidorhynchus corniculatus (No. 684).—Milson Island, Hawkes- 
bury River, May, 1915 (with Halteridium). 

Ailuroedus smithi (No. 720).—Bunya Mountains, Q’land, Oct., 
1919 (a few large spherical Leucocytozoa). 


13. Haematozoa not Detected. 
ieee humeralis (No. 26).—Stradbroke Island, Q’land, Sept., 
1919 


Eudyptula minor (No. 62).—Encounter Bay, Feb., 1921. 
Hieracidea occidentalis (No. 260).—Narrabri, Jan., 1918. 


118 . 


Calyptorhynchus leachi (No. 289).—Narrabri, Nov., 1916 (2); 
Dorrigo, Jan., 1918. 

Cacatua. gymnopis (No. 293).—Beltana, Aug., 1921. 

Platycercus pennanti (No. 304). —Mount Wilson, June, 1915; 
Bunya Mountains, Q’land, Oct., 1919. 

Platycercus eximius (No. 311). —Dubbo, July, 1915. 

EKurystomus pacificus (No. 341).—Scone, ‘Oct., 1917. 

Cacomantis flabelliformis (No. 362). —Milson Island, Hawkesbury 
River, Jan., 1915 (young bird); Stradbroke Island, Q’land, 
Sept., 1919. 

Petrochelidon nigricans (No. 386).—Stradbroke Island, Q’land, 
Sept., 1919. 

Orthonyx spinicaudus (No. 464).—Dorrigo, Jan., 1918. 

pro iucle lunulata (No. 488).—Bunya Moutains, Qland, Oct., 

Chthonicola sagittata (No. 501).—Baan Baa, Jan., 1917. 

passers uno Reet (No. 509).—Baan Baa, Jan., 1917 ; Dubbo, 

uly, 

Acanthiza lineata (No. 511).—Bunya Mountains, Q’land, Oct., 1919. 

Lope pusilla (No. 512).—Bunya Mountains, Q’land, Ook. 


Sericornis frontalis (No. 519).—Canobolas, Oct., 1916 (2 birds); 
Mount Irvine, June, 1915. 

eases cyanochlamys (No. 530a).—Bunya Mountains, Q’land, 
Cb; 


_Malurus melanocephalus (No. 542).—Mummulgum, near Casino, 


Dec., 1916. 
peas leucogaster (No. 559).—Stradbroke Island, Q’land, Sept., 
1919. 


Colluricincla rufigaster (No. 573).—Stradbroke Island, Q’land, Sept., 
1919. 

Climacteris leucophaea (No. 593).—Bunya Mountains, Q’land, 
Oct., 1919. 

Zosterops dorsalis (No. 599).—Bunya Mountains, Q’land, Oct., 1919. 

Stigmatops ocularis (No. 639).—Stradbroke Island, Q'land, ‘Sept., 
1919. 


Anthochaera carunculata (No. 675).—Scone, April, 1917. 

Sericulus chrysocephalus (No. 726). —Mummuleum, Dec., 1916. 
Corvus coronoides (No. 732).—Bumberry, Jan. 1916. 

Corvus cecilae (No. 733).—Stradbroke island, Q’land, Sept., 1919. 
Strepera graculina (No. 735).—Scone, April, 1917 (2 birds). 
Passer domesticus (Sparrow). —Sydney, June, 1917. 


Dea 


THE EXTERNAL CHARACTERS OF POUCH EMBRYOS OF 
MARSUPIALS. 


No. 4. ~PSEUDOCHIROPS DAHLI. 


By Freprric Woop Jones, D.Sc., F.Z.S., 
Professor of Anatomy in the University of Adelaide. 


[Read June 8, 1922.] 


Pruate VI. 


4 


For all the pouch embryos of this interesting form I am 
indebted to the authorities of the Perth Museum. The 
animal was first described by Professor Collett in 1895. In 
1915 it was placed by Matschie in the sub-genus Pseudo- 
chirops, when that author split up the large Genus 
Pseudochirus of Ogilby. Pseudochirops dahli and P. archeri 
are the only Australian members of the sub-genus, the 
other seven constituent species being confined to New 
Guinea. From the external characters of the pouch embryo 
it would appear to be a particularly interesting form, and 
one that is undergoing remodelling in response to the 
demands of a comparatively recent radiation. 


Har.—Hair is first visible in the 80 mm. stage, at which 
time the embryo is flesh coloured. The 50 mm. embryo 
shows no trace of body hair, though the specialized tactile 

_ vibriscae are present. When the embryo has reached 105 mm. 
the body is entirely clothed with short hair, the general 
colour of which is light brown. The skin of the embryo is 
free of pigment. 


Hair Tracts.—The hair tracts are charted from male B, 
Perth Museum, the embryo, which is shown at pl. vi., being 
105 mm. in total length. Upon the head are numerous 

definite hair fields arranged in a rather complicated manner 
(see fig. 1). 

(A) Immediately behind the naked rhinarium a field 
of short hair shows a uniform forward direction; the free 
tips of the short hairs extend to the superior margin of the 
naked rhinarium, and to the upper margina of the narial slit. 

(B) Behind this is an area extending backwards to the 
anterior angle of the eye, and laterally downwards to the 
mysticial region. In this field the hair is directed forwards 
and towards the mid-line, so that the areas of the two sides 
of the snout meet in the mid-line at a hair ridge. This field 


field (C) streams backwards away from the forwardly- 


120 


is also marked off by a definite ridge from the field imme- 
diately in front of it. , 

(C) Above and around the eye, the hair streams upwards 
and backwards so that it leaves a well-marked divergent 
parting above and in front of the orbit where the hair of 


( 


Hair tracts of the head (from Specimen Male B, 
Perth Museum, 105 mm.). 


directed hair in field (B). 

The area (C) meets its fellow of the opposite side in the — 
mid-line of the head and ends behind at a convergent hair- 
line which runs roughly from the crown of the head to the 


‘peel oy} 07 esopo Yo yno se poyuesorded st opoline otf], 
‘(mUL GOT ‘wnesny, 449d ‘q aTeyy woutoedg) syoesy Ae Py 
: 


121 


122 


posterior margin of the palpebral fissure. At the crown of 
the head a whorl (V in fig. 1) is developed upon each side 
of the middle line at the upper end of this convergent line. 


- 


ws 


~ Hig: 3. 
i aeial vibri iscae (from Specimen A, Perth 
Lo Museum, 80 mm.), — 
The next tract (D) is a complex one, for radiating from 
a single mid-line whorl (W in fig. 1) situated upon the dorsal 
surface of the head opposite the margin of the ears, the hair 


Wig. 4. 
Brachial vibriscae (from Specimen Female A, Perth 
Museum, 80 mm.). 


streams in three different directions: (1) forwards and 
downwards, where it meets (C) at the convergent hair-line ; 
(2) outwards to clothe the dorsal and posterior surface of 


123 


_the ear; and (3) backwards and downwards Inte the general 


body stream. 
Upon the side of the face the ace field, . which 
starts at the lower narial ‘Margin and turns backwards below 


Bie, Dd. 
Calcaneal vibriscae (from Specimen Female A, 
Perth Museum, 80 mm.). 


(A) and (B), becomes continuous with the sub-occular field. 
The hair in this tract (E) is directed backwards and 
slightly downwards. At the angle of the mouth: it joins with 


is. 6: 
Rhinarium (from Specimen Female A, 
on sass U a mm. ) 


the » pack paialty doped ee of the eee jee These 
combined backwardly-directed streams meet the pre-auricular 
part of the field (D) and continue the convergent hair-line, 


124 


which, starting at the crown of the head, ran past the 
posterior angle of the eye to the lower jaw near its angle. 

The hair tracts of the body and limbs need little 
description to supplement their diagrammatic representation 
in fig. 2. 


Bip. -7. 
Form of the external ear. 
A, 35 mm. stage. 3B, 50 mm. stage. 


WY 


ANY iti 
wy : 


= 


Wis. 8. 
Form of the external ear. 
C, 80 mm. stage. D, 105 mm. stage. 


_ There are no hair reversals upon the body or limbs, and 
no whorls, crests, or partings are present. The main stream- 
lines are caudad and ventrad on the body and ventrad and 
post-axial on the limbs. 


Hts 


125 


Hair is continued to the ungual extremity of the 
phalanges of both manus and pes; the heels in the fully- 
haired embryo are almost wholly naked. 

The hair when first present is so pale as to be practically 
colourless; when the embryo is fully haired the hair is of a 
very pale brown. 


Fig. 9. 
Left manus, 35 mm. stage. 


Sensory Papillae and Vibriscae.—Sensory papillae are 
developed at the 35 mm. stage and vibriscae are present at 
50 mm. The first papilla to appear is the ulnar-carpal. 

Facial Vibriscae—The mysticial set consists of 6 rows 
of papillae (in Collett’s description 7), giving rise to 2, 5, 7, 6, 
6, and 5 backwardly-directed, pale vibriscae, respectively. 
The supraorbital papilla is large, and gives origin to 2 
vibriscae. The genal bears 6 long sensory hairs. The 
interramal is inconspicuous, with 2. pale hairs; and the 


126 


submental consists of small papillae with rather trivial but 

early developed hairs (see fig. .3). ‘etsy ee 
Brachial Vibriscae.—The ulnar-carpal papilla. is: large, 

and gives rise to a brush of half a dozen or so pale bristles. 


Di 


) 


WN 


pC eS Re 
Ea) 
Lies ly 


Re== 
a ° 


— 


ce Press 
2 
<= 


*. 


Fig. 10. 
Left manus, 105 mm. stage. 


The anconeal and the medial brachial give rise to a single 
hair each (see fig. 4). 

Crural Vibriscae.—The crural papilla is well developed. 
Two stout tactile hairs arise from it, one of these bristles 
being, in all specimens, considerably longer than the other 
(see fig. 5). } 


127 


~The Rhinarium.—The rhinarium is roughly triangular 
in shape and distinctly grooved in the middle line. The 
surface’ is finely granular. The narial slits are bounded 
above entirely by naked skin, but their lower margins are 


Fig. 11. 
Left pes, 35 mm. stage. 


pubescent behind. The infranarial portion of the rhinarium 
runs to the upper lip, forming a very definite portion of its 
medial area. (see fig. 6). | 

The External Har (see figs. 7 and 8).—In all stages 
which I have examined: the auricle has been folded back- 
wards. This is true of the 35 mm. embryo. The whole 
process of the development of the pinna may be described 
as a progressive simplification. Two well processi antihelicis 
appear, but only one persists as a meatal operculum. A 
_well-marked bursa in.the 80 mm. embryo becomes reduced 
to an insignificant depression in the 105 mm. stage. Of the 


128 


tragus and antitragus, the tragus alone persists in any degree 
of finished development. 
The Manus (see figs. 9 and 10).—The digital formula of 


—» 


LT Vie 
oes 


} 


= 


i 
1 
is 


~ 
WN 


| iN) R\2 ‘ | ' 
: \\\| wid. Wy yes Te ~ 
W. | 
AN 


ail 


~ & <a; 
ya , 


Ah, 
LOM /, 
SS 


Fig. 12. 
Left pes, 105 mm. stage. 


the haired embryo is 3>4>2>5>1. In the earlier stages 
the 4th digit is longer than the 3rd. In the 80 mm. embryo 
there is a definite tendency for the digits 1 and 2 to stand 


129 


in opposition to digits 3, 4, and 5; but by the 105 mm. stage 
this dual division of the manus has ceased to be at all well 
marked. Herein lies the great interest of the manus of this 
form. Apical pads are present on all digits and are striated. 
Interdigital pads are striated and are 3 in number, interdigital 
pad i being fused with the thenar pad. 


: 
\ 


ar 


Mize 
“ie 
re 
ii 
my 
Heh 
[¥ 
SS 


i 
YER 
/d TB 
_ ees 
aie 


Ss 


thy 
mn 
Ue 
IT i 
size 
7 
MM) 


Fig. 13. 
Three stages in the development of the pad at the base 
of the first pedal or digit. — 
A,50mm. B,80mm._ C, 105 mm. 


The Pes (see figs. 11, 12, 13).—The digital formula is 


—_— 
4>5>2,3>1; the syndactylous toilet digits being relatively 
far longer in the early stages. Apical pads are present and 
striated. Interdigital pads striated, but the striations are 


a 


130 


somewhat ill defined in older embryos. The point of‘ out- 
standing interest is the fusion of interdigital pad 1 with the 
thenar pad, and the progressive diminution of the striations. 
Between the 80 mm. and the 105 mm. stages considerable 
readjustment takes place in the disposition of the sole in the 
region of the ‘base of the first pedal digit (see (B) and (C) in 
fig. 13). Professor Collett has described in an embryo “about 
100 mm., from snout to vent’’ 2 pads at the base of the big 
toe, but gives no description of the pads in the adult. This 
change is presumably to be correlated with the loss of 
opposibility of digits 1 and 2 to digits 3, 4, and 5 in the 
manus. It would seem that the animal had somewhat fallen 
from the arboreal standards of its immediate stock. The 
diagnosis made from the conditions of the hands and feet is 
borne out by Professor Collett’s account of its habits: “During 
the day time it hides amongst the colossal boulders, and 
leaves the rocks only at night, when it ascends the trees in 
search of food’’ (P.Z.S., 1897, p. 332). » 

External Genitalia.—The pouch is normal. The opening 
directed cephelad, and 4 mammary areas are present. 


DESCRIPTION OF PLATE VI. 
Pseudochirops dahli. 


Pouch young photographed against a background of +,-1n. squares. 
Specimen Male B, Perth Museum, 105 mm. 


131 


THE TERTIARY BROWN-COAL BEARING BEDS 


OF MOORLANDS. 


By Str Doucias Mawson, D.Sc., B.E., and Freperick 


Cuapman, A.L.S., F.R.M.S. 


[Read June 8, 1922.] 


Page 
I. InTRODUCTION Bs oe ce Be % ae lS 
Il. GenerRaAL PHyYSIOGRAPHY AND GEOLOGICAL FEATURES 132 
Ill. THe Tertrary Srrata at Mooranps ... & sae cael 5 151 
Division 1.—Recent Surface Formation. 
Division aes ea (Lower Pliocene) Oyster 

Bed. 

Division 3.—Janjukian (Miocene) Marine Beds. 

Section A.—Green and _ yellow 
Clays, Marls, and Sands. 

Section B.—Uight-grey and dark- 
grey Clayey Marls and Cal- 
careous Muds. 

Section C.—Marine Limestone 
and Carbonaceous Muds usually 
pyritised. 

Division 4.—Janjukian Lacustrine Carbonaceous 
Beds with Lignite. 
* IV. ComMPaRISON BETWEEN THE BEDS IN SouTH AUSTRALIA 
ied ee A she oat LAG 


AND VICTORIA 


I. INTRODUCTION. 


The occurrence of brown coal in Tertiary strata in the 
vicinity of Moorlands, a railway station on the Pinnaroo 
line about 87 miles from Adelaide, has led to very consider- 
able mining activity thereabouts during the past two and a 


half years. 


As a result, much valuable geological informa- 


tion has been collected in an area where otherwise no 
geological section of the beds would be available. 

We are particularly indebted to Mr. A. C. Broughton, 
the representative on the field of the principal mining com- 
pany, for assistance in procuring data and material amplify- 
ing such as was secured on our own visits, which date back 
to the inception of the present mining enterprise. The 
Government Geologist has also favoured us with information 
required relating to the Government bores. 

Though the main bulk of these notes were prepared 
more than two years ago, publication has been delayed in 
ease important additional information relating to the beds 
should accrue as a result of mining development. In the mean- 


132 


time much has been made public in the Mining Reviews | 
of the Department of Mines under reports by the Govern- 
ment Geologist, Mr. L. K. Ward; the Chief Inspector of 
Mines, Mr. L. J. Winton; and the Engineer for Boring, 
Mr. C. F. Duffield. A short note has also been contributed 
by Mr. A. C. Broughton.) 
The scope of this present paper is accordingly restricted 
to generalized notes upon the strata, more particularly a 
correlation with the Tertiary beds of other localities. 


Il. GENERAL PHYSIOGRAPHIC AND GEOLOGICAL FEATURES. 


Moorlands is situated on a nearly level mallee-covered 
plain which extends from the Murray River (some 10 miles 
to the west) eastward into Victoria. Over all this area 
undulations of the surface are rarely conspicuous. Perhaps 
the most noteworthy of such is the long, low rise known as 
Marmon Jabuk Range, which trends in a general N.N.E. 
and §.8.W. direction across the country just to the north 
of the Moorlands coal field. Such rises are often composed 
of flexed Tertiary beds, but, at other times, much more 
ancient rocks come to the surface in these more highly 
elevated portions. The latter are frequently slaty beds not 
unlike certain of the ‘‘Adelaide Series,’’ and probably of 
late pre-Cambrian age.) At times more highly altered 
sedimentary rocks appear; for example, a strongly developed 
chlorite schist was entered in a well sunk about one mile 
south-east. of Moorlands railway station. Ancient igneous 
rocks are, probably, not uncommon underlying the Murray 
mallee lands, as evidenced by the outcrops of pink granite at 
Mannum, at Murray Bridge, and to the south of Coonalpyn ; 
also, the appearance of a broad intrusive sheet of gabbroic 
rock, now much modified by age, exposed in the railway 
cutting, on the line to Moorlands, about two miles beyond 
Tailem Bend. . 

But, though there is unquestionably a considerable 
diversity in the underlying strata, the surface features of 
these mallee plains, as a rule, give little indication thereof, 
for there is developed everywhere at the surface a hard 
travertine formation which varies from a few inches to a 
few feet in thickness. It is thickest where it overlies Tertiary 
strata and thinner where the more ancient rocks underlie it. 


(1)See Mining Review, Nos. 13, p. 21; 32, pp. 32-38; 33, pp. 
64-78; 34, pp. 31, 32, 34-39, 43-50; 35, pp. 25, 26, 28-42, 47-55. 
(2) ‘“‘Notes on the Geology of the Moorlands (South Australia) 


Brown Coal Deposits,’ by A. C. Broughton, Trans. Roy. Soc. 
S. Austr., vol. xlv., 1921, pp. 248-253. 


(3) Vide Paper read by T. W. E. David, Trans. Roy Soc. 
S. Austr., vol. xlvi., Nov., 1921. 


133 


Where one has to cross this country in a vehicle, the traver- 
tine, outcropping in knobby and platy masses, is, for the 
most part, developed uncomfortably close to the surface, but 
in depressed areas it is usually covered by a thin mantle of 
sand or sandy soil. Occasionally, superficial sand is heaped 
up into low dunes, which aid to modify the monotonous level 
of the country. 

Even in the areas occupied by them, it is a rare thing 

to locate the Tertiary beds definitely by the discovery of 
fossil remains at the surfaces, though some of the larger 
molluscan remains have been found amongst the surface 
travertine in specially favoured spots. Bores put down in 
search of brown coal are gradually furnishing definite data 
as to the distribution and details of deposition of these beds ; 
but so far, beyond the fact that some part of the trans- 
Murray mallee country is underlain by Tertiary formations 
and some is not, little absolutely definite is known. 

The probability is that only in minor areas does the 
ancient primary rock come to the surface. Elsewhere fossili- 
ferous Tertiary beds, in greater or less thickness, either 
horizontal or but slightly inclined, are to be expected as the 
uppermost formation, but owing to the semi-arid climate are 
masked at the surface by the development of a dense super- 
ficial layer of travertine or aeolian sand formation. 

The steep cliff-like banks of the lower Murray river, 
which latter approaches within 10 miles of Moorlands, furnish 
good geological sections through marine Miocene (Janjukian) 
beds which have been long explored. Patches of lower 
Pliocene (Kalimnan) limestone are also dispersed in this region, 
above the Janjukian, but are less regular than the latter. 

To the east of the river, in the vicinity of Tailem Bend, 
these beds thin out very quickly, and within a few miles of 
the river the older formation comes to 'the surface over 
considerable areas. In this neighbourhood only scattered 
shallow pockets of the Tertiary, principally Kalimnan, are 
met with. This condition persists eastwards until Moorlands 
station is closely approached, when a decided and continuous 
low dip to the east carries the pre-Tertiary rocks downwards, 
so that an ever-increasing thickening of the Tertiary beds 
is met with as the Victorian border is approached. This state 
of affairs is illustrated by the data from various bores quoted 
by Mr. L. K. Ward.“ Whereas ‘‘bed rock’’ (pre-Tertiary) 
is encountered at depths ranging between 50 ft. and 100 ft. 
in most of the areas where mining activity is now proceeding 
at Moorlands, a bore sunk at a point 40 miles to the east 
penetrated 852 ft. of Tertiary strata before meeting the 
older bed rock. 


_@ Min, Rev., No. 38, pp. 72-74. 


134 

Recent mining exploitation has shown that the brown 
coal seams are developed at or near the bottom of the 
Tertiary formation. As a consequence of the dip to the east, 
the brown coal’ formation comes to the surface, or nearly 
approaches‘ it, in the Moorlands area. It is this fact that 
has led commercial exploitation to especially favour this par- 
ticular locality, for open-cut mining is thus made, possible, as 
opposed to the more expensive method of winning the coal 
by deep mining, entailing additional costs in labour, pump- 
ing, and’ timbering. 

In connection with mining exploration, bores are now 
being sunk ‘at intervals of 300 yards, and even closer, in 
places. This close boring is steadily accumulating a fund 
of information invaluable for discussion of the contour of the 
surface upon which the Tertiary strata was laid down. Thus, 
also, will much light be shed upon the question of erosion 
intervals, if such do actually exist between the beds of the 
Tertiary strata. But until all the bores can be referred to 
the same datum level, which has not so far been done, final 
statements in regard to the above must be deferred. 

In Mr.: Broughton’s paper ) reference is made to one 
line of bores which he had related to the same datum level 
by means of a dumpy-level traverse. As a result, he shows 
that the floor of the Tertiary formation is slightly undulating 
and the coal beds occupy the depressions in this old land 
surface of low relief. On this evidence it is assumed that 
the shallow basins containing coal are isolated by rises in 
the floor of ancient slaty rocks. 

These depressed basins may have been ponded areas in 
the coal-forming period, where plant life thrived and was 
preserved in sodden beds. Subsequent marine sedimentation 
overlapped the lignite-filled basins and extended as a con- 
tinuous shéet over much of the former old land surface. 

The very unequal thicknesses of brown coal met with in 
the various bores is accounted for, at least partly, by such 
an original ’ accumulation in basins. But it yet remains to 
be shown to what extent irregularities in the coal beds are 
due to wash-outs of the nature “of erosion by contemporaneous 
streams .of ‘the coal-forming period, which tracks would be 
afterwards obliterated by silts rendered highly carbonaceous 
from ligneous matter transported from erosion areas else- 
where. This and the question of a general erosion interval 
at the upper limit of the lignite beds, with its bearing also 
upon the extent and distribution of the residual lignite are 
matters to be settled when the boring operations are com- 
pleted. '- , 


(5) Loc. cit. 


135 
Ii]. Toe TERTIARY Strata at MooRLanps.. 


Surface sand. 
Travertine. 
Pale-yellow sand. 


Hard limestone 
_ (oyster. bed). 


Division 1. 
Recent. 
Division 2. 
Lower Pliocene - 
(Kalimnan). 


f 


-.' Seetion: A. 


=o oi © 8m 


a 
Division 3. = Sa 
Miocene Marine .- = See Sachin B 
(Janjukian). = | '|E= ses 3 seep ig 
Light-grey to dark- 
grey clayey marls and 
calcareous muds. 


Section. C. 
Dark-grey _pyritic 
sandy limestone above. 
. Dark-grey carbona- 
ceous pyritised muds 
‘below. 
Lignite with clay. 


Lignite. 


Division 4. 
Miocene Lacustrine. 


gece Genk 


“ 


G. 


.. Lignite with clay. 


%-10 


] 


+h. 


100-ft. level. Clay with’ lignite. 


Pre-Cambrian. 


Fig, L. 

The geological section, set forth herewith, represents the 
Tertiary strata existing in the vicinity of the main shaft sunk 
by the Murray Coal and Oil Co., which is at the same time the 
vicinity of Government Bore 25. It is the. particular locality 
where most of our detailed observations were made.': In order 
to better illustrate the average nature of the. beds, a slight 

generalization has been assumed. 


136 


In the following descriptions, the major formations are 
taken in descending order from the surface. 


Division 7. 
RECENT SURFACE FORMATION. 


Travertine, with more or less blown sand, forms a surface 
layer. The travertine is the usual surface form developed 


under the conditions of a semi-arid climate. In some parts | 


of the field, the travertine sits directly upon the Ostrea lime- 
stone, and is little more than an addition to and modification 
of it. Elsewhere, sandy beds intervene which may or may 
not belong to the marine series “below. 
Division: 2. 
A KALIMNAN (LOWER PLIOCENE) OYSTER BED. 


This is.a hard marine limestone, usually buff-coloured. 
This rock is what may be termed a ‘‘ragstone,’’ or roughly 
fracturing limestone, filled with oyster shells and pectens. 
It would appear from the results of borings that this bed is 
not continuous over the field. In many cases, however, it 
is undoubtedly included with the surface travertine in the 
one entry. The fossils determined from this horizon are: — 


Pelecypoda—Ostrea aremcola, Tate, a smooth upper 
valve; O. sturtiana, Tate, probably the commonest 
fossil in this bed ; Pecten antiaustralis, Tate, rare; 
P. palmipes, Tate, a fragment only ; Spondylus 
arenrcola, Tate, a restricted Aldingan species, rare. 

Pisces—Isurus hastalis, Agassiz, sp. (tooth). This fossil 
is rather worn, but the outline leaves no doubt-of its 
identity. A common Victorian Kalimnan fossil. 


Embedded in this limestone small pebbles of white quartz 
and other rocks are occasionally met with. Of the latter 
the following were collected:—A water-worn pebble, 3 in. 
in length, of a rock resembling a mica granulite; several 
chips of slate up to 2 in. in length; and two pieces of basalt, 
one water-worn, the other partly faceted. 


The latter basalt specimen measures about 4 in. by 2 in. 
It is a grey rock with open steam holes, the vesicles being 
drawn out by flow. In microscopic section, laths of labra- 
dorite felspar are noted to be the dominant feature. A flow 
arrangement of the felspars around the steam holes is evident. 
A large corroded fragment of plagioclase exhibiting poly- 
‘synthetic twinning is also to be seen. A small amount of 
interstitial pyroxene is still visible. A considerable quantity 
of secondary serpentine is present and appears to be chiefly 


137 


after olivine. Magnetite is present in moderate quantity, 
and also leucoxene. Traces of limonite and haematite are 
also present. This rock is not similar to any specimens of 
the Mount Gambier basalt which we have at hand for com- 
parison, but in general character it is like some of the Mel- 
bourne basalts.. The question that arises is from what locality 
and by what means did it become transported to its present 
situation ? 
Division 3. 
JANJUKIAN (MIOCENE) MARINE BEDS. 


Section A. 
GREEN AND YELLOW CLAYS AND SANDS. 


These are soft clayey and sandy beds, often notably 
calcareous, of greenish or buff colour. Yellowish and reddish * 
mottlings and streaks may appear where these beds rise 
above ground water level, thus exposing the iron content to 
oxidising influences. ; 

‘In some portions of the field borings have revealed strata 
in this section of the beds of great uniformity. In such cases 
a general buff colour is assumed. The greenish tint due to 
glauconite granules, which is an outstanding feature in other 
areas, is, in these situations, largel¥ suppressed. In all cases, 
however, at least a little glauconite can be detected, on close 
examination. The buff-coloured silt forming the bulk of such 
beds is exceedingly fine-grained and of low specific gravity. 
The average grain size amongst the observable discrete par- 
ticles in one of the finer bands proved to be 1/100 of a 
millimetre diameter. The coarser particles are well rounded 
except for very minute flecks of mica. The sand grains of 
the. coarser beds of this series are unusually rounded and 
polished. In fact, a large part of these buff-coloured sedi- 
ments is loessial in character. Such beds are particularly 
well represented in a bore at the cross roads, some two miles 
north of Moorlands station. The buff-coloured component 
of this sediment possibly originated as wind-blown dust from 
the interior of the continent, picked up and transported by 
the ancestral Murray-Darling River system. Judging from 
the highly glauconitic character of some of the beds, such were 
probably laid down in current-disturbed waters at a con- 
siderable depth. 

Fossils are reasonably common only in the more highly 
glauconitic portions of this section. One such bed of a bright 
apple-green colour and of a calcareous nature was examined 
in detail for fossils. Polyzoa were found to be comparatively 
numerous and rotaline foraminifera not uncommon. Besides 


138 


these there are occasional Brachiopods, Bivalves, joints of 
Alcyonarians, and ossicles and spines of Echinoderms. 


A similar bed was struck in several of the Victorian — 
mallee bores, where it was seen to be a bed of glauconitic 
and shelly sand and glauconitic chert. 


A small series of fossils selected from a highly glaheotitas 
bed of this division yielded the following : — | 

Foraminifera—Truncatulina ungeriana, d’Orb, sp. 
Fairly abundant. 

Anthozoa—Mopsea tenisom, Chapman. The smaller and 
slenderer joints of this coral are very abundant in 
the washings. 

Echinodermata—Cidaroid spines, various; (?) Antedon, 
ossicle. 7 ; 

Polyzoa—Cellaria, sp.; Adeona obliqua, MacGillivray ; 
Porina gracilis, Milne-Edwards, sp.; Steganoporella 
patula, Waters, sp. ; Cellepora gambierensis, Busk ; 
Retepora permunita, MacGillivray. 

Brachiopoda— Terebratulina — catinuliniformis,. Tate; 
Magasella woodsiana, Tate. 

Pelecypoda—Pecten foulcheri, T. Woods (a fragment). 

Pisces—The following fish teeth were collected on the 
field and handed to the authors, it being understood that all 
were obtained from Division 2 (the Miocene) of the section, 
and principally, if not entirely, from Section A. They are, 
in nearly all cases, much rolled and often imperfect. The 
determined species are :— 

Odontaspis elegans, Agassiz, sp.—This widely distributed 
species is here quite abundant, being represented by about a 
dozen specimens. It is more commonly met with in the 
European Miocene and Pliocene than in Australia, and we 
have never before seen so many specimens of this form from one 
locality. It occurs in the Eocene and Lower Miocene.of New 
Zealand, and in the Miocene and Lower Pliocene of Victoria. 

Odontasms incurva, Davis, sp.—This species is here repre- 
sented by one specimen, nearly perfect, but for the left side 
of the base being missing. It is fairly common in the 
Miocene and Lower Pliocene of Victoria, and it has a much 
more extensive range in New Zealand, where it begins its 
history in the Upper Cretaceous (Danian) and ranges up to 
the Lower Miocene. 

Odontasms exigua, Davis.—This species is here repre- 
sented by several examples. It is quite a new record for the 
Australian Tertiary deposits, as hitherto it has only been 
found in the New Zealand Lower Miocene of the Trelissick 


139 


Basin. The widely expanded base, relatively small size, and 
the short stout fang are distinguishing characters. 

Odontaspis attenuata, Davis, sp.—Two imperfect examples 
are referred to the above species. It was recorded from the 
Oamaru Series of New Zealand. In Victoria it is found in 
the Miocene, and is of the same age in South Australia 
(Aldingan Series). It also occurs in the Lower Pliocene 
(Kalimnan) at Beaumaris, Port Philip. 


Section B. 
LIGHT-GREY AND DARK-GREY CLAYS. 


These underlie the greenish and yellowish beds of the 
previous section. Faint bluish and chocolate colours some- 
times appear. Sandy beds are not uncommon. These beds 
are fairly rich in calcium carbonate. Pyrites may enter in 
noticeable quantity at the base of this section. The pre- 
dominating dark colour is due to carbonaceous matter, and 
is doubtless derived from beds of the underlying lignite where 
exposed in other areas undergoing erosion at the time of 
deposition of these clays. 

Apart from the colouration due to carbonaceous matter, 
the beds of this section are very similar to the previous, and 
appear to be merely a continuation downwards of the same 
general type of sedimentation. Small marine fossils, prin- 
cipally gasteropods, are sparsely distributed in this section. 


Section C. 


MARINE LIMESTONE AND CARBONACEOUS MUDS, USUALLY 
PYRITISED. 


This section immediately overlies the lignite series. It 
is never very thick. A hard dark-grey or, less frequently, 
light-yellow sandy limestone, full of marine fossil remains, 
usually limits it above and rests upon a dark sandy mud full 
of molluscan and other remains. Both the limestone and the 
mud are usually highly pyritised. At its base it often shows 
. distinct brecciation, including fragments of slate. So that 
in all probability there was in progress in the vicinity, at the 
time of the accumulation of this bed, a great deal of erosion. 
The black carbonaceous and pyritous condition of the sedi- 
ments points to material worn down from the previously 
formed lignitic bed. 

It is without doubt equivalent to Tate’s horizons‘) in 
the Croydon Bore at 1,376 ft. (‘‘Bituminous clay and black 
sand ; Turritelia aldingae’’), and 1,681 ft. (‘‘Bituminous shale ; 


(6) Trans. Roy. Soc. S. Austr., vol. xxii., 1898, p. 195_ 


140 


casts of gasteropods in chalcedony, calcite and iron pyrites, 
some shell matter; Twurrtella aldingae, Mesalia stylacris, 
Fibularia gregata, Cellepora’’ ). 

Examined as to its nature, the following details are 
observable : — 

When moist this bed is somewhat sticky in texture, but 
easily washes down, on account of the large proportion of 
fine quartz sand which it contains. 

Fine Washings.—These consist largely of quartz grains, 
sub-angular to rounded, with usually a very high polish. 
They were undoubtedly of aeolian origin and carried into 
the marine deposit either by estuarine river flows or high 
winds, but most probably through the former. The quartz 
grains measure about ‘4 to 1 mm. in diameter. Admixed with 
the quartz grains are numerous granules of pyrites and frag- 
ments of polyzoa and calcareous molluscan shells; some of the 
shell fragments show attached crystals of iron pyrites. 


Coarser Washings.—About one-sixth of the material 
remains as coarse washings and contains by far the larger 
number of fossils. These are more or less fragmentary, or 
often partially pyritised. Some of the smooth, rounded, 
white, quartz pebbles are beautifully coated by a superficial 
deposit of brassy pyrites laid on in electroplate fashion. 

The fossils determined from this horizon are: — 

Anthozoa—Mopsea tenisonn, Chapman; IM. hamilton, 

_ Thomson, sp. 
Echinodermata—-A cidaroid spine, cf. Leocidaris. 
Polyzoa—Cellepora gambierensis Busk; Adeona obliqua, 
MacGillivray; Porina gracilis, Milne- Edwards, en 
Brachiopoda—Terebratula tateana, T. Woods. 
Pelecypoda—Limopsis insolita, Sowerby, ss Arca 
(Plagiarca) cainozoica, Tate, sp.; A. (Barbatia), 
sp.; Crassatellites pee Tate, sp.; Cardita 
latissima, Tate; C. delicatula, Tate; Cardium 
monilitectum, Tate; Corbula pyxidata, Tate; C. 
ephamilla, Tate. 

Gasteropoda—-Collomia parvula, Tate; Huspira effusa, 

Tate; Turritella aldingae, Tate; TJ. trestera, 
Tate; 7. platysmra, T. Woods; Diala, sp.; Letorium 
oligostirum, Tate, sp.; Marginella, cf. strombiformis, 
T. Woods; Scaphella pagodoides, Tate, sp.; S. 
pueblensis, Pritchard, sp.; Ancilla ligata, Tate, sp.; 
Turris, sp.; Actaeon, cf. subscalatus, Cossmann. 


Pisces—-Fish otolith (Teleostean). 


141 


Division 4. 
JANJUKIAN LACUSTRINE, CARBONACEOUS BEDS WITH LIGNITE. 


These are fresh water lacustrine beds of very variable 
thickness which rest, more or less horizontally, upon the up- 
turned edges of steeply inclined slates and other strata of 
Lower Cambrian or earlier age, and are overlain by the 
Janjukian marine beds. There appears to be some evidence 
of erosion of the upper limits of the ligneous series, but the 
.extent of such will be better defined as exploration of the 
field proceeds. 

It is evident, however, that the age of the fresh water 
lignite-bearing beds is either Miocene or pre-Miocene. It is 
most reasonable to regard these beds as having formed con- 
temporaneously with either one or other of the strongly 
developed Tertiary lignites of the adjacent State, Victoria. 
Referring to these latter occurrences, the Altona lignite series 
is intercalated with typical marine Oligocene (Balcombian) 
fossil bands. But the Morwell beds, so far as can be judged, 
occupy a former lacustrine or coastal swamp area, of which 
the hinterland river deposits of the Miocene Dargo High 
Plains form a part. Judging from the palaeogeography of 
the region, therefore, the Morwell lignite is Miocene in age. 
The reference of the carbonaceous lacustrine beds at Moor- 
lands to the Morwell (Miocene) period is indicated by the 
fact that they appear to be comparable with a similar forma- 
tion, underlying the polyzoal rock in the mallee, and which, 
besides containing lignite, includes typical J anjukian marine 
shells, T'rigonia lamareki, found occurring at the top of the 
J anjukian, and the foraminifer, Cyclammina complanata, 
Chapman, the latter eae found at the base of the J anjukian 
at Anglesea. 

Ligneous beds of a character similar to these at Moor- 
lands are now being actively explored at three other localities 
in South Australia, namely, at Clinton, on the west side of 
Gulf St. Vincent, almost at its northern extremity; at Hope 
Valley, one of the northern suburban areas of Adelaide; 
and at Noarlunga,(% 23 miles south of Adelaide, adjacent to 
Gulf St. Vincent. Furthermore, similar beds were encoun- 
tered in a Government bore put down at Bower, 20 miles 
west of Morgan, several years ago. There is also other 
evidence from boring operations in South Australia, indicating 

(7) Vide Dept. of Mines, S. Austr.;. Min. Rev., Nos. 34, pp. 

29-31, 40, 51; 35, pp. 26, 27, 43-46, 55-56. 


(8) Ninas Bai. Nos. 14, p. 10; 20, pp. 19, 40, 41; 33, pp. 25-37; 
34, pp. 27-29, aa Al, 42; 35 pp. 13-15. 


(9) Min. Rew. No. 33, pps (8,79: 
(10) Min. ee Nos. 20, p. 11;..28._ pp..26-28;: 29. p. 23. 


142 


that prior to the period of formation of the Miocene marine 
beds there was a great and widespread development of Tertiary 


MORGAN 
e 
RS 
G 
7 PORT WAKEFIELD 
?e 
Zz 
» 
re) 
FOR 
ADELAIDE 
z (\ 
a 
Ss j HOPE VALLEY 
. ADELAIDE 
fee 
3 
£* 
6 
® NOARALUNGA 
a) 7 
3 ATAILEM BEND 
% J BMOORLANDS 
% LAKE 
cee ALEXANDRINA ‘ 
SUE 
Ss oo" : 
1 % Murrd AWE 
eee’ 
LoS Ene 
& sare a at ht, a 
2 


Fig. 2. aS 
Map illustrating South Australian Brown Coal Occurrences. 


tresh water ligneous strata. This fortunate circumstance 
has bequeathed to the present day large reserves of brown coal. 


143 


The South Australian areas where lignite beds have been 
proved belong to one or other of two main regions of Tertiary 
sedimentation. Of these, one is Gulf St. Vincent and its 
borderlands; to this the Clinton, Paradise, and Noarlunga 
occurrences are to be referred: ‘The other is the great Murray 
Gulf, so marked a feature of Tertiary times in Australia, 
when a shallow sea extended all over the south-east lands of 
South Australia, part of Victoria, and even reaching well 
into New South Wales territory. Bower and Moorlands 
belong here. The published results of the Government bores 
indicate that the lignite beds of the Gulf St. Vincent area 
are embedded in, and entombed beneath, strata of a more 
sandy nature than is the case on the Murray side, where very 
fine clayey and calcareous silts are a greater feature. This 
is no doubt attributable to muddy waters delivered to the 
latter area by the great ancestor of the present Murray- 
Darling system. 

In this connection there is some evidence indicating 
that, in all probability, the brown coal of the Gulf St. 
Vincent region may reach a greater degree of freedom 
from ash than that influenced by the muddy waters of the 
ancestral Murray. So far as at present explored, this is the 
case, as can be noted by a comparison of the seams at Moor- 
lands with those at Clinton. The Mines Department Records 
give composite analyses‘) of all lignite beds met in all bores 
to date, respectively, in each of thes fields as follows : — 


Moorlands. Clinton. 

Moisture at 105° C. -..: Dileat 51°47 

Volatile matter a 21°38 24°29 

Fixed carbon ... e 13°82 15°16 

Ash oe ie? oy pease ao 9°08 

100°00 100°00 

Sulphur content ea 2°66 2°01 
Average moisture after 

alr-drying... ae 15°74 16°94 


In arriving at these figures the Government Geologist, 
Mr. L. K. Ward, differentiates the carbonaceous beds as 


- follows :— 


“‘Lignite’’ is such as contains not less than 24 per cent. 
of ash. 

“Lignite with clay’’ is the designation for beds contain- 
ing between 24 and 50 per cent. ash. 

“Clay with lignite’’ distinguishes beds containing over 

50 per cent. ash. a 


(11) Telnaes Bores 1 to 34, at Moorlands; and 1 to 4, at 
Clinton. 


144 


Though the above figures represent the average case, so 
far as exploration has yet proceeded, seams are met with of 
much better quality than the average here stated. For in- 
stance, at Moorlands the better quality lignite is illustrated 
in Bores 23, 24, and 31 by seams, respectively, 13 ft., 18 ft., 
and 16 ft. in thickness, averaging only about 7 per cent. ash. 
At Clinton, in Bore 1, a band of lignite, 15 ft. in thickness, | 
containing less than 6 per cent. ash, is recorded; and in 
Bore 2 there is a 15-ft. seam with 7°3 per cent. ash. 


“The ultimate relative merits of the two fields, from an 
economic standpoint, depend upon factors far more important 
than slight differences in composition. The shallow depth of 
the formation, the more water-tight nature of the strata, 
the comparative freedom from serious quantities of ground- 
water, are all factors which favour the Moorlands area. On 
the other hand, convenience in geographic situation for 
marketing the fuel is favourable to the Gulf St. Vincent 
localities. 


It is probable that, in the aggregate, great quantities 
of brown coal will be proved, eventually, at the base of the 
Tertiary formations in South Australia. This is most re- 
assuring from an economic standpoint, for the mineral fuel 
resources of the State are otherwise limited to the semi- 
bituminous, Triassic(2) coal of the small Leigh Creek basin. 


The best of the Leigh Creek seams (5) averages about 
20 ft. of coal carrying 18 per cent. ash and 20 per cent. of 
moisture displaceable at 105° C. Of this seam the best 6 ft. 
averages as follows :— 


Per cent. 
Moisture lost at 105°C. ... Js so ee 
Volatile matter oh be: viel  . oe 
Fixed carbon sp he a ih eee 
Ash ae ae wee ey se 6°9 
100°0 


This appears to be the best that the Leigh Creek semi- 
bituminous coal field can produce as a working proposition. 
So taken into consideration with remoteness from centres of 


population, it is obvious that the Tertiary brown-coal forma- . — 


tions are likely to be the State’s mainstay in the matter of 
fuel resources. 


(12)The occurrence of Macrotaeniopteris wianamattae and 
Thinnfeldia odontopteroides in these beds fixes the age as Triassic. 


(13) Vide Government Geologist’s Report, Min. Rev., No. 27. 


145 


In general appearance and composition, the Moorlands 
lignite is very similar to the better known Victorian occur- 
-rences—for example, that of Morwell. The latter, however, 
is freer from ash than any of the South Australian so far 
examined. 


No fossil remains other than plants have yet been 
detected in the lignite and associated clays of Division 4 
in the Moorlands section. The plant remains in the lignite 
itself are singularly well preserved, often retaining the woody 
structures so as to be clearly distinguishable in the hand 
specimen. 

The coal cuts like cheese and, consequently, is easily 
excavated. In the seam, or when freshly mined, it is of a 
_ black or deep chocolate-brown colour, but exposure to the 
air at the surface soon causes it to dry out, accompanied by 
contraction which develops shrinkage cracks, eventually lead- 
ing to a crumbling of the mass. As drying proceeds the 
general colour becomes lighter until it is literally a ‘‘brown 
coal.’’ 

The remains of small trees, sticks, and reeds are clearly 
distinguishable as blacker lignite embedded in a browner base. 
The latter is observed to be constituted principally of the 
remains of leaves and small twigs. Some of these leaves are 
very fresh and tough, so that they may be picked out in 
large pieces or even entire. They are so thin and translucent 
as to be capable of mounting on glass slips as transparencies. 
Selected examples, thus mounted, have been submitted to an 
authority on Tertiary floras, Mr. Henry Deane, M.A., F.L.S., 
who comments upon them as follow :—‘‘Broader-toothed leaves 
evidently Proteaceous, but not ‘Banksia.’ Venation resembles 
more nearly that of a short leaf among my specimens of 
Telopea speciosissima. Specimens of Lomatia Fraseri and 
L. wheifolia have been compared, but in these the dentation 
of the margin is invariably too strong. The small narrow 
leaf bears a strong resemblance to some leaves with entire 
margins of Banksia marginata.”’ 


Embedded at random in the leafy base are particles of 
two varieties of resin. The more abundant is very dark 
coloured, practically black. It occurs characteristically in 
large, elongated, tear-like drops, usually about 1 cm. in 
diameter, but often considerably larger. In its general 
appearance and occurrence it is akin to certain of the grass 
tree gums of to-day, but darker in colour, which, however, 
is a feature which would be expected to develop with age. 


In considerably less quantity and in smaller particles 
of an irregular shape, usually about one-third of a centimetre 


146 


or less, is a light yellow-brown resin. Like the former, it 
is peilinnt on fracture face but dull externally. 


As to the question whether the lignite has grown where 
it is now accumulated, considerable evidence of growth im situ 
has been observed. | 


_ Rootlets have been distinctly noted traversing the inter- 
bedded lignitic clays. A small stump, about 8 in. across, 
was got in these beds when excavating the original shaft on 
Mineral Lease 1233, Hundred of Sherlock. Furthermore, 
the arrangement of the components of the lignitic mass is 
adverse to any suggestion of water transportation. For 
example, the particles of resin are embedded at random, and 
there is no evidence of water sorting as between the leaf 
remains and the massive woody tissues. | 


IV. CoMPARISON BETWEEN THE BEDS IN SoutH AUSTRALIA 
AND VICTORIA. 


The Tertiary deposits of the old Murray Gulf appear to 
extend continuously from the eastern limits of the Mount 
Lofty Ranges across into the mallee of Victoria. The strata 
at Moorlands represent depositions on the western side of the 
old gulf, and a study of the beds is of especial interest for 
comparison with those on the Victorian side, which latter have 
already been written upon in regard to the deep borings in 
the mallee. (4) 


In the present collection of fossils by far the larger 
number of marine shells show an aspect comparable to the 
Aldingan fauna of Professor Tate, both as regards the Lower 
(Miocene) series and the Upper (Lower Pliocene) beds. On 
the Victorian side the same deposits, both lower and upper, 
show the middle and upper part of the Hamiltonian facies,(® 
as proved in the deep borings of the mallee. Thus, in the 
latter locality there were no shells of Spondylus arenicola, 
Pecten palmipes, and P. consobrinus, although others were 
common to both localities. This dissimilarity in faunas, so 
closely adjacent, would suggest a bar or rocky ridge separ- 
ating the sea of that period. The curious deepening of the 
bathymetrical sedimentary conditions, shown in the Tin- 
tinara Bore, have been referred to as due to trough folding: 


(44) Chapman, F.: Cainozoic Geology of the Mallee and 
other Victorian Bores, Rec. Geol. Surv. Vict., vol. iii., pt. 4, 1916. 


(15) A regional word, here coined: to express the combined 
faunas of the lower and upper. Muddy Creek beds with the inter- 
calated limestone of the Grange Burn, ranging from the Bal- 
combian to the Kalimnan. 


147 


or even to the formation of a _ rift-valley with infilled 
material. (16) 


This present series affords data which help one to divide 
the Miocene (Janjukian) beds of both the old Murray and 
the Spencer Gulfs into three sections, as exemplified here and 
in the Victorian bores of the mallee area :— 

(3) Glauconitic bed, yellow clays and sands. 

(2) Polyzoal rock, or grey to whitish calcareous sand, 
passing downwards into a pyritous and quartzose 
deposit, with marine fossils. 


(1) Carbonaceous beds with lignite. 


For the present we may regard these as equivalent to the 
Upper, Middle, and Lower Miocene, respectively, although 
their limits are not clearly marked. 


It is interesting to note here that the order of the Beds 
2 and 3 are reversed, as far as lithology goes, at Torquay 
(Spring Creek beds) ; but it must also be borne in mind that 
the molluscan facies from the two glauconitic series would 
eed differ. 


(16) Chapman, F.: Ree. Geol. Surv. Vict., vol. iii., pt. 4, 
1916, p. 407. 


148 


CONTRIBUTIONS TO THE ORCHIDOLOGY OF AUSTRALIA 
AND NEW ZEALAND. 


By R. 8S. Rocers, M.A., M.D. 
[Read July 13, 1922.] 


I. ADDITIONS. 


Diuris brevifolia, 0. sp. Planta gracilis, glabra, cir- 
citer 15-40 cm. alta. Folia 4-8, linearia v. setacea, acum- 
inata, non torta, erecta, 7-12 cm. longa. Flores 1-4, laxe 
racemosi, lutei cum notationibus bruneis paucis. Sepalum 
dorsale ovatum, recurvum, circiter 15 mm. longum; sepala 
lateralia herbacea, linearia, acuminata, parallela, patentia. 
Petala breviter petiolata, circiter 14 mm. longa, lamina 
elliptica. Labellum sepalo dorsali aequale aut paulo longiore ; 
lobus intermedius rhombo-cuneatus, basi intus, carina ~ 
duplici, lateralibus plus quam duplo longior. 

Slender, glabrous; leaves generally 4-8, linear or 
setaceous, acuminate, not. twisted, very erect, rarely reaching 
beyond the middle of the stem. Flowers solitary, or in a loose 
raceme of 2-4, on slender pedicels, yellow with a few brown 
markings, much smaller than those of D. sulphurea. Dorsal 
sepal ovate, recurved; yellow with two dark-brown spots on 
the dorsum, one on each side of the base. Lateral sepals 
much longer, green, linear, parallel, spreading below the 
labellum or slightly recurved. Petals nearly as long as the 
lateral sepals, shortly stalked, spreading or recurved; lamina 
a canary-yellow, elliptical, about 11 mm. long. Labellum 
yellow, at least as long as the dorsal sepal and generally 
longer; lateral lobes less than half as long as the central 
lobe, generally about 5 mm., not very wide, margins entire, 
tips recurved; middle lobe rhombo-cuneate with» depressed 
antero-lateral margins; lamina with two closely approximated 
parallel raised lines on the basal half continuous with the 
anterior central keel, the lines surrounded by a conspicuous 
dark-brown border. Anther without a definite point, rather 
higher than the viscid disk of the rostellum. Lateral 
appendages of the column subulate or linear-falcate, about 
the same height as the viscid disk. 7 

South Australia: Longwood and other parts of the Mount 
Lofty Range; Myponga; Mount Compass; Port Elliot; 
Kangaroo Island. November-December. 

This plant. has long been confused with D. suwlphuwrea, 
which it superficially resembles, but from which it differs 


149 


in its short setaceous and relatively numerous leaves; in its 
much smaller flowers and in its labellum, which is at least 
as long as the dorsal sepal and bears two raised longitudinal 
lines. . 

Its relation to other South Australian members of the 
genus is indicated in the following table: — 


Flowers not blotched or spotted on their upper 
surface, but of a uniform colour. 
Flowers purple or heliotrope (drying yellowish- 
brown); lateral sepals greatly es 


petals, ‘about 5 em. long .. D. punctata 
Flowers canary-yellow ; lateral sepals only 
slightly exceeding petals 4 D. pedunculata 


Flowers yellow with conspicuous dark-brown or 
purple-brown markings or blotches. 
Lateral lobes of labellum large, as long or 
nearly as long as middle one. 
Lateral sepals greatly exceeding petals in 
length, often nearly twice as long; leaves 
6 or more, setaceous or almost so ... ... D. palustris 
Lateral sepals shorter than, or approxi- 
mating in length to the ‘petals: leaves 
not setaceous. 
Lateral sepals crossed; blotches generally 
distinctly demarcated from the yellow 
ground-colour; 2 Cricaee raised 
lines at base of labellum... ... .... .... D. maculata 
Lateral sepals nearly parallel ; flowers 
wall-flower colour, dark blotches merg- 
ing into yellow ground-colour; 1 raised 
line at base of labellum ... .. D. longifolia 
Lateral lobes of labellum very much shorter 
than the middle one. 
Two raised longitudinal lines along base of 
labellum. 
Flowers with small dots and short linear 
markings; leaves linear and rather lax, — 
often 17 cm. long ... D. palachila 
Flowers with 2 conspicuous ‘brown dots at — 
base of dorsal sepal and conspicuous 
oblong brown border round the raised 
lines; leaves usually more than 5, 
setaceous or nearly so; _ short (about 
7 or 8 cm.) and very erect ... D. brevifolia 
One longitudinal raised line along base of 
labellum. 
Flowers with similar markings to D. 
brevifolia,; and in addition a brown 
transverse blotch near the tip of middle 
lobe of labellum; leaves usually 2, 
rarely 3, long, lax, linear «.... ... ... «D. sulphurea 


Prasophyllum Brainei, 0. sp. Planta viridis, gracilis, 
12-24 cm. alta. Lamina folii basi dilatata, deinde anguste 
linearis vel setacea, spicae circiter aequalis. Spica laxa, 5-10 


150 


cm. longa. Flores 10-24, sessiles, virides, pediculus per- 
brevis. Segmenta perianthii glandulosa. Sepalum dorsale 
erectum v. recurvum, ovato-lanceolatum, concavum, acumina- 
tum, circiter 5°75 mm. longum; lateralia libera, patentia, 
leviter apicibus recurva, circiter 7 mm. longa. Petala erecta, 
anguste oblonga v. lineari-lanceolata, circiter 5 mm. longa. 
Labellum sessile, basi subgibbosum, ad columnam erectum ; 
deinde recurvum sigmoideum, ad apicem contractum et 
brevissime ciliatum, marginibus crenulatis; pars callosa ovato- 
lanceolata, marginibus anticis ciliatis, paulum ultra primum 
flexum producta ; pars membranacea latiuscula, alba, inferiore 
dimidio levis, in superiore dimidio rugosa. Columnae laciniae 
late oblongae; apicibus obtusis obliquis; rostellum in 
altidudine excedentes. Anthera rostello brevior. 

A slender green plant, 12-24 cm. high. Lamina of leaf 
dilated at the base, thereafter setaceous or narrowly linear, 
about same length as the spike. Flowers almost sessile in 
a loose spike, their very short pedicel subtended by a broad 
short blunt bract, entirely green, recurved from the axis of 
inflorescence; ovary 4-5 mm. long, obovate; all segments very 
glandular. Dorsal sepal erect or recurved, concave, ovate- 
lanceolate, acuminate, slightly contracted at the base. Lateral 
sepals quite free, spreading but only slightly divergent, tips 
slightly recurved, concave on upper surface, narrow lanceolate. 
Lateral petals erect, narrowly oblong or linear-lanceolate, not 
very acute. Labellum sessile; rather gibbous at the base, 
but not protruding between the sepals; proximal part erect 
against the column, the margins entire until a little beyond 
the middle; thereafter recurved so as to form a complete 
sigmoid flexure, laterally contracted towards the apex, the 
margins crenulate and very shortly ciliate from the first 
bend to the extreme tip; the callous portion dark green, 
ovate-lanceolate, extending from the base to a little beyond 
the first flexure, its termination concealed by the lateral con- 
traction of the lamina at that point, its margins shortly 
ciliate; the membranous part rather wide, whitish, smooth 
in the erect part, rather rugose anteriorly, very elandular, 
more or less tomentose towards the tip. Lateral appendages 
of the column relatively large, broadly oblong, with blunt 
oblique tips, basal lobes distinct, exceeding the rostellum in 
height. Anther shorter than the rostellum. 

Named after Mr. A. B. Braine, an ardent collector and 
student of Victorian orchids. 

Victoria: Ringwood (E. E. Pescott). October. 

The new species approaches the green forms of P. fuscum, 
but materially differs from it in the much less complicated 
structure of the labellum and shorter lateral appendages of 
the latter. 


151 


Pterostylis humilis, 1. sp. Planta robusta, perbrevis, 
2-3. cm. alta. Folia 4-6, rosulata v. subrosulata, sessilia, 
imbricata, 0°5-2°5 ecm. longa, ovata v. oblonga. Flos unicus, 
sessilis; ovarium basibus foliorum in parte obtectum. Sepalum 
' dorsale ovato-lanceolatum, circiter 15 mm. longum cum petalis 
connivens. Galea subangusta, ap ce acutiuscula. Labium 
inferius oblongo-cuneatum, erectum, sinu acutissimo, lobi 
subulati circiter 13 mm. longi galeam multo superantes. 
Labellum unguiculatum, lineari-oblongum, ad apicem ob- 
tusum sensim contractum; lamina circiter 11 mm. longum, 
linea longitudinalis elevata in medio; appendix linearis, 
eurvata, penicillata. Columna circiter 10 mm. longa; anthera 
terminalis, obtusa, bilocularis, erectiuscula; lobi supeviores 
laciniarum breves lineares, inferiores longi falcati acutissimi. 
Stigma perprominens, infra columnam mediam, late cordatum, 
lobis distinctissimis. 

A rather stout plant of low stature, arising from two 
more or less conical or globose tubers. Leaves (in the flower- 
ing stage) 4-6, rosulate or subrosulate, sessile, sheathing, 
imbricate ; lamina of varying length, oblong, ovate or oblong- 
ovate. Flower solitary apparently sessile, the ovary partly 
hidden by the sheathing bases of the leaves. Dorsal sepal 
ovate-lanceolate, about 15 mm. long (when extended), con- 
nivent with the petals to form a rather narrow erect galea, 
apex of galea rather acute but not prolonged into a filiform 
point. The base of the lower lip oblong-cuneate, erect; lobes 
subulate (hardly filiform), including a very acute sinus, 
embracing the galea. Labellum reddish-brown, on a movable 
irritable claw, oblong-linear, tapering a little towards a very 
blunt and slightly recurved tip; lamina traversed by a raised 
longitudinal line with a corresponding groove below; basal 
appendage linear, curved, penicillate. Column (extended) 
about 10 mm. long. Anther terminal, bilocular, quite blunt, 
rather erect. Wings of column with a short linear upper 
lobe or tooth ; the lower lobe long, falcate, very acute. Stigma 
very prominent, situated below the middle of the column; its 
two lobes very distinct, together forming a broadly cordate 
disk. Rostellum linear-oblong situated between the bases of 
the anther loculi and connected to the stigma by a split tube. 

New Zealand: The Haunted Whare, near Waimarino 
(H. B. Matthews). | aa 

_ Mr. Matthews states that his specimens were removed 
from their natural habitat near the base of Ruapehu (within 
three miles of perpetual snow), and cultivated in Auckland, 
200 miles north of their native locality. He thinks that the 
change to abnormal conditions may have produced a dwarfed 
growth in the plant. Along with his spirit specimens, he 


152 


forwarded a photograph of a fruiting specimen. This indi- 
cates a plant of different habit, with a stature of 11 cm.; 
with leaves on well-marked petioles and lamina from 3°75-6 cm. 
long. It is probable that the scape becomes elongated after 
pollination, so as to assist in the maturation of the fruit, as 
happens in the case of many Australian orchids, notably in 
the genus Corysanthes. On the other hand, it must be remem- 
bered, that in certain other species of the genus, dwarfed 
specimens are by no means infrequent. This is particularly - 
true of P. cucullata, where dwarf forms are often to be found 
growing side by side with normal plants. These show such 
a departure from the type that even experienced botanists 
like Sir J. D. Hooker and Robt. Brown fell into error and 
described them as separate species. 


Mr. Matthews further states that unlike other members 

of the genus, the flower is reversed, the labellum being upper- 
most, owing apparently to a retroflexion of the column on 
the ovary. 
. The new species appears to correspond rather closely to 
the description of P. trifolia, published by Colenso in New 
Zealand Inst., xxxi. (1898), 281. As only a single specimen 
of Colenso’s plant was discovered, and that is not available 
for comparison, it is not possible to say whether the two 
orchids are identical. Cheeseman, however, regards P. trafolia 
as conspecific with P. venosa, which differs from Mr. Mat- 
thews’ plant in column and in some other respects. 


Caladenia pumila, n. sp. Planta pumilissima, per- 
hirsuta, 5-10 cm. alta. Folium basi-amplexicaule, circiter 
3-6 cm. longum, hirsutissimum, lineare v. olbongo-lanceo- 
latum. Caulis robustiusculus, hirsutissimus. Flos solitarius, 
albus, comparate pergrandis. Segmenta perianthii sub- 
aequalia, paene glabra, acuminata, non-caudata. Sepalum 
dorsale lanceolatum, concavum, erectum, incurvatum, circiter 
2°5 cm. longum, basi latiusculum; lateralia latiora, libera, 
patentia, lanceolata. Petala patentia sepalis angustiora. 
Labellum breviter unguiculatum, ovatum, apice obtusum, 
obscure 3-lobatum, circiter 15 mm. longum 11 mm. latumque; 
dimidio inferiore ad columnam erectum marginibus in- 
tegerrimis, deinde recurvum marginibus carneis serrulatis 
v. crenulatis; lamina transverse complanata, callis carneis 
anguste linearibus 4-6 seriatis prope medio terminantibus. 
Columna circiter 13 mm. longa, incurvata, in dimidio 
superiore late membranaceo-dilatata. Anthera incumbens, 
valvata, bilocularis. Pollinia 4, typica. ; 

A very hairy species of low stature. Leaf relatively large, 
linear or oblong-lanceolate, clasping at the base; stem rather 


153 


stout, with a rather large free acute bract close to that 
subtending the terminal pedicel. Flower solitary, white, 
relatively very large. Segments of perianth white, usually 
without markings but sometimes with a faint pink stripe on 
the outside; nearly equal in length, glabrous except at the 
extreme base, not contracted into caudae, gradually diminish- 
ing into finely acuminate non-clavate points, the latter rarely 
glandular. Dorsal sepal erect, lanceolate, incurved, concave, 
about 2°5 cm. long, the base rather wide. Lateral sepals 
wider, free, spreading, lanceolate, somewhat contracted at 
the base. Petals spreading, lanceolate, narrower than the 
lateral sepals. Labellum on a short claw, white with narrow 
pink margins, a few pink splashes on the lateral lobes, 
obscurely 3-lobed, ovate, blunt at the apex, about 15 mm. 
long and 11 mm. wide; the lower half erect against the 
column with entire margins; thereafter recurved with serru- 
late or crenulate margins; the lamina flattened transversely, 
the calli pink narrowly linear in 4-6 rows ending near the 
middle. The column nearly as long as the labellum, incurved, 
speckled with pink, widely winged in its upper half. Anther 
shortly mucronate, valvate, 2-celled. Stigma circular, with 
short triangular rostellum in its upper border between the 
bases of the anther cells. Pollinia in 2 pairs, of the usual 
type. Ovary exceedingly glandular-hairy. 

Victor: Bannockburn (EK. E. Pescott). September- 
October. 

The new species differs from C. Patersoni in its dwarfed 
habit, in the absence of tentacles to the perianth segments, 
and in the absence of definite glandular tips to those seg- 
ments. Its sepals are about equal in length to the petals, 
whereas they are considerably longer than the latter in C. 
Patersom. The tip of the labellum is blunt and the margins 
practically entire in C. pumila, whereas the tip is acute and 
the margins acutely toothed in the other species. 


Prasophyllum Frenchii, F. v. M., var. Tadgellianum, 
n. var. Flowers pale greenish-yellow ; or yellow with chocolate 
markings down the middle of the perianth segments, also down 
the middle and on the sides of the labellum. Lateral sepals 
connate. 
é Victoria: Mount Hotham (5,100 ft.); Mr. A. J. Tadgell. 
December, 1914. 
_ New South Wales: Mount Kosciusko (7,300 ft.).; Dr. 
' Green. December, 1921. 
The specimens from these two alpine localities would 
appear to be morphologically identical. In coloration they 
E 


154 


differ from F. v. M.’s plant, and also from each other. The | 
chocolate markings on the Mount Kosciusko specimens are an 
exceedingly conspicuous feature and give to the flower a very 


distinctive appearance. Fortunately a single bloom was suffi- | 


ciently fresh to enable this observation to be made. The 
Mount Hotham plants were dry, and like many prasophyllums © 
in that condition very difficult to examine. All the specimens | 
differed from the type by the possession of connate sepals. | 
The union or otherwise of these segments is a variable feature 


in P. Suttonu, another alpine member of the genus, and was § 


not considered sufficiently important to indicate a specific — 


difference. It is possible, however, that closer acquaintance | 


with this plant may cause it to be given the higher rank. 


Prasophyllum australe, Br., var. viscidum, n. var | 
Plant slender; flowers rather smaller than the type, dark red | 
or prune coloured, with many darker blotches or spots, very 
viscid. y 

Victoria: Alberton, Gippsland; ‘‘in sandy soil, swampy | 
in winter time’; Mr. A. J. Tadgell. January(?), 1921. | 

Of this very interesting and unusual variety, Mr. Tadgell — 
writes : —‘‘It is so viscid, that it is quite a trouble to detach | 
it from the drying-sheet. . . . It is scarred like a leper, | 
on flowers and stem.’’ 


Caladenia carnea, Br., var. aurantiaca, n var. Very — 
slender, about 14 cm. high. Flowers 1 or 2, the second one 
on a filamentous pedicel. Perianth segments white on the 
inside, striped with green on the outside. Labellum pure 
white with exception of the tip and the calli, which are deep 
orange in colour; calli in 2 rows, with large clavate heads 
and slender stalks ;.tip entire, its margins without denticula- 
tions or calli. 


Victoria: Alberton, Gippsland; A. J. Tadgell. October, 
1920.4 «: 
The contrasting colours of this dainty little Caladenia 
give it a very characteristic appearance and charm. There 
are no transverse stripes on the lamina as in the type. The 
stem and ovary are distinctly hairy; the leaf narrow-linear 
and almost glabrous. 
_ 2. Norss. 


DENDROBIUM DicuPpHUM, F. v. M. Leaves 4, lanceolate, 
with 5 prominent nerves, 15-18 cm. long and about 2 cm. wide. 
‘Flowers white with purple centre, 9-12, in a raceme on a 
slender peduncle about 36-40 cm. long. Perianth segments 


155 


longitudinally veined. Sepals similar, acute, oblong-elliptical, 
about 16 mm. long and 5 mm. wide. Petals obovate, about 
18 mm. long and 10 mm. wide. Spur 2-fid; the lower seg- 
ment oblong-cylindrical, very obtuse, about 3 mm. long. 
Labellum about 13 mm. long and 10 mm. wide (flattened out) ; 
| 3-fid; middle lobe obtusely oblong; lateral lobes wide and 
rounded ; lamina with rather numerous calli distributed along 
the nerves, but chiefly in about 6 rows terminating near the 
centre, longitudinally veined. Northern Australia: Groote 
Eylandt, Gulf of Carpentaria; Mr. N. B. Tindale. August, 
1921. 
SPIRANTHES AUSTRALIS, Lindl. Column erect, about 
3 mm. long, fleshy, contracted in its lower half, clinandrium 
dilated ; anther valvate, 2-celled, blunt or minutely apiculate, 
inclined against the back of the stigma and reaching to about 
its upper border; the wings represented by a membrane on 
each side stretching between the ‘‘filament’’ of the anther and 
the stigmatic-plate, adnate to the pedicel (“‘style’’) of the 
latter and also to the lateral margins of the stigma itself, 
forming a pouch between the male and female elements of 
the column, the bases of the pollinia released quite early 
from the anther so as to rest in the bottom of this pouch. 
Stigmatic surface U-shaped, large and slightly sloping down- 
wards. The rostellum (including the disk) almost equal in 
length to the stigmatic surface, arising from the upper border 
of the latter, forming a long narrow membranous structure 
much exceeding the anther in height; with a short rigid 
acute bifid base persisting after the membranous portion has 
been removed, or as a long split membranous structure after 
removal of the disk and pollinia only. The disk slate 
coloured, long narrow elliptical (or “‘boat-shaped’’) accom- 
modated in a fork of the rostellum and covered by a mem- 
branous capsule derived from the latter and containing a 
viscid fluid; the capsule attached to the lateral margins of 
the disk and traversed by a central vertical furrow. Pollinia 
lamellated, in 2 pairs, the latter club-shaped or pyriform, 
granular; the apices of the pairs lightly united, exposed above 
the anther-case and attached by a short caudicle to the 
back of the disk; capsule of disk easily ruptured artificially 
so as to permit removal of the pollinarium. 


The structure of the column in Spiranthes australis is 
comparable to that in the genus Prasophyllum, but in the 
former the wings of the anther-filament are adnate not only 
to the pedicel of the stigmatic-plate, but also to the margins 
of the stigma itself. 


ad Rogers, Trans. Roy. Soc. S. Austr., xly. (1921), p. 264. 
E 


156 


To rupture the capsule of the disk artificially, a light 
but appreciable force is required. An attempt to produce 
rupture by 15 minutes’ exposure to chloroform vapour, as 
in Darwin’s experiment, was unsuccessful. The facility with 
which the whole pollinarium may be removed is strongly sug- 
gestive of an insect-pollinated plant, yet the examination of — 
large numbers of plants revealed the pollinia still am situ and 
in only one instance was pollen found adhering to the stigma. — 
No difficulty was experienced in removing the pollinia from 
fully expanded flowers, although R. D. Fitzgerald states that 


this is impossible. This botanist writes: —‘‘I could discover ~ 


no trace of a rostellum or disc of any kind. In this flower 
the persistence with which the pollinia remained behind the 
stigma, though left naked by the shrinking back of the anther, 
is very peculiar. No transfer of the substance of the stigma 
on the point of a pin or a bristle induces them after opening 
of the flower to come forth for the chance fertilization of 
another flower. It even requires some violence to break them 
as the more friable portions turn towards the anther’ (Aus- @ 
tralian Orchids, vol. i.). q 


In the numerous specimens which I have examined from — 
South Australia, Victoria, and Queensland, the ‘“‘split © 
rostellum,”’ the ‘‘boat-formed disk,’ the ‘‘easily-removable — 
pollinia’”’ of Darwin were all present. In fact, Darwin’s 
description of these structures in S. autumnalis may be 
accepted as a most accurate description of the same structures 
in the Australian species. The capsule of the disk does not 
appear to split so readily as in the European species. In no 
other respect does it appear necessary to modify the great 
observer’s classical description. 

It can hardly be doubted that Fitzgerald’s observations 
were conducted on a species with which Australian botanists 
are unfamiliar, a species which, so far as is known to the 
writer, is unrepresented in our national collections. 

Whether Smranthes australis is self- pollinated or other- 
wise is a matter which cannot yet be.regarded as settled. 
Undoubtedly a large number of seed capsules are frequently 
to be found on some spikes, whereas spikes from other locali- 
ties display comparatively few. 

CaLOCHILUS PALUDOSUS, R. Br. Victoria: Bayswater; 
Mrs. Edith Coleman. 23/10/21. 


THELYMITRA MEGCALYPTRA, Fitzg. Victoria: Grampian | 


Mountains;- J. W. Audas. 31/10/20. 


T. Macmixuani, F. v. M. This species, which is usually 
salmon coloured, has been collected by Mr. E. E. Pescott at 
Bannockburn, Victoria, of a deep rose or crimson colour. 


157 


T. LonGiFoLta, Forst. South Australia: Mount Pata- 
warta (3,060 ft. elevation), 365 miles north of Adelaide; Mr. 
B. B. Beck. 5/10/20. 

This represents not only the highest elevation, but also 
the furthest north at which any orchid has been recorded in 
this State. A few other orchids from the same locality are 
noted below. 

T. GRANDIFLORA, Fitzg. Victoria: Nar Nar Goon; J. W. 
Audas. 18/10/20. 

T. URNALIS, Fitzg. South Austraha: Bugle Ranges; 
National Park; Dr. and Mrs. Rogers. October, 1921. 

This orchid was described in 1882, but has never until 
this season (1921) been reported since its discovery. They 
correspond in every respect to Fitzgerald’s description and 
illustration, except that the tooth in front of the column is 
not always present. The plants were not numerous and were 
found growing alongside of 7. antenmfera, Hook., and 7’. 
rubra, Fitzg. The flowers are yellow and bear dark-brown 
stripes on the outer sides of the sepals identical in appearance 
to those occurring in 7’. antennifera. It is quite possible 
that the plant may be a hybrid between this and the other 
species mentioned above. 

Diuris aurea, Sm. WNew South Wales: Barrington 
Tops (5,100 ft.), near Patterson; Mr. A. N. Burns. 14/12/21. 

D. LonerFotia, Br. With two well-developed parallel 
raised lines on the lamina of the labellum. The lamina 
normally bears only one such line. This fact is rather im- 
portant, because Bentham makes the number of such lines 
a prominent feature in the classification of the members of 
this genus. Mr. Max Jacob, who collected these specimens 
at Cherry Gardens in this State, informs me that they are 
by no means uncommon this season (1921) in that locality. 

PRASOPHYLLUM SUTTONII, Rogers and Rees. In addition 
to Buffalo Plateau (Victoria), where this alpine species was 
discovered, it has reached me from the following localities : — 

New South Wales: Mount Kosciusko (7,300 ft.); Mr. R. 
Helms. February. 

Victoria: Baw Baw Mountains (5,060 ft.) ; Mr. C. French, 
jun., January; Mount Feathertop (5,000 ft.); A. J. Tadgell, 
December. 

Tasmama: Summit of Ben Lomond (5,000 ft.); A. 
Simson. March. 

Further acquaintance with this orchid shows that the 
lateral sepals are not always free, but are subject to varia- 
tion, as in certain other members of the genus. 


158 


P. austRALE, Br. In this species, as in P. elatum, the 
lateral sepals are very consistently connate. A departure from 
this rule is noted in the case of certain specimens collected 
by Messrs. E. E. Pescott and C. French, jun., at Monomeith, 
Victoria, among which there are a number of flowers with 
five sepals. 


P. BREVILABRE, Hook. f. Vzctorra: Mount St. Bernard 
(4,000 ft.) and Mount Hotham (5,000 ft.), Australian Alps; 
Mr. A. J. Tadgell. December, 1913. : 


PTEROSTYLIS PYRAMIDALIS, Lindl. Western Austraha: 
Jarnadup; Miss Knox-Peden. 1/9/21. 


All of these specimens are unusually tall, some attaining 
a height of 33 cm.; the plant quite slender and erect. Leaves 
at the base 3 or 4, ovate, acute, not strictly rosulate, shortly 
petiolate or clasping; passing into leaf-like sessile alternate 
bracts, ovate to lanceolate in shape, diminishing from below 
upwards, the lower ones generally with serrulate or crenulate 
margins, sometimes 16 in number. Flower much larger than 
that of P. nana; the galea from base to crest often nearly 
2 cm. long. The inturned tooth between the lobes of the 
lower lip appears to be invariably present, but in other 
respects the general habit of the plant is very different from 
that of P. nana. | 


P. cYcCNocEPHALA, Fitzg. Wew South Wales: Mount 
Kosciusko (7,300 ft.); Dr. Green. 29/12/21. 


P. pEepoGLossa, Fitzg. Tasmania: Brown _ Mountain, 
Port Arthur; Miss A. L. Rogers. 2/5/19. 


P. MitcHevyui, Lindl. South Awstralia: Mount Pata- 
warta (3,060 ft., 355 miles north of Adelaide); Mrs. R. 8. 
Rogers. 14/10/15. 


P. rura, Br. South Austraha: Mount Patawarta 
(3,060 ft., 355 miles north of Adelaide); Mr. B. B. Beck. 
5/10/20. 

Labellum on a very wide and elastically membranous claw, 
longer and narrower than usual, very hairy, the hairs of 
exceptional length. 


CoRYSANTHES, sp.(?) Capsules dehiscing, on slender 
pedicels, 13-15 cm. long. The remains of the flower enabled 
me to identify the genus but not the species. The plants 
were sent from a Victorian locality, and are of interest in 
showing how the pedicel, which is almost sessile during the 
flowering season, becomes enormously elongated in order that 
the seed capsule-may receive the benefit of wind and sun in 
the process of maturation. Corysanthes blooms in June or 


159 
July, and the specimens were collected by Mrs. Edith Cole- 
man in December. Members of the genus propagate chiefly 
by the vegetative method, and such specimens as these are 
rarely found. 

CALADENIA DILATATA, Br. South Australia: Mount Pata- 
warta (3,050 ft.); Mr. B. B. Beck. 5/10/20. 

C. GLADIOLATA, Rogers. South Australia: Cherry Gardens; 
Mr. Max Jacob. 26/9/21. 

This plant was discovered at Hornsdale, 175 miles north 
of Adelaide. It has not hitherto been recorded from any 
other habitat. 

CuILocLoTTis Gunnir, Lindl. New South Wales: Mount 
Kosciusko (7,300 ft.); Dr. Green. 29/12/21. Victoria: 
Australian Alps (Mount Hotham 6,000 ft., and Mount St. 
Bernard 5,100 ft.); Mr. A. J. Tadgell. December, 1913. 


160 


THE PHYSIOGRAPHY OF THE MEADOWS VALLEY, 
MOUNT LOFTY RANGES. 


By E. O. Tearez, D.Sc. 3 
(Communicated by Professor W. Howchin.) 


[Read July 13, 1922.] 


The broad outlines of the development of the physio- 
graplical features of South Australia have been admirably 
traced out by Prof. Walter Howchin.@ Relics of ancient 
topography, and the deposits of ‘‘dead rivers’? have been 
widely recognized, and in piecing this evideice together, the 
great importance of tectonic movement in the form of warping 
and faulting has been rightly emphasized, for it has certainly 
had an important influence in the development of the existing 
conditions of climate and topography. The highlands of the 
Mount Lofty Ranges provide a noteworthy example in this 
direction. They were recognized by Howchin as owing their 
origin to block faulting, whereby several segments moved 
differentially with regard to each other. 

Remnants of very ancient and mature topography are 
still to be found alongside of fresh and youthful features, 
where erosional activities, revived by differential earth move- 


ments, are energetically working towards the destruction of 


those relics which throw so much light on the past geographical 
history of the region. 

The observations of this paper centre around the Meadows 
Valley, in the southern portion of the highlands, and were 
gathered by the writer) during his geological and soil survey 
of the Kuitpo Forest Reserves and their vicinity. 

The nearest part of this region to Adelaide lies about 
20 miles in a straight line to the S.S.E., and the valley trends 
in a S.S.W. direction for 10 or 12 miles, eventually joining 
the Finniss River through a narrower and more steeply graded 
course. It is a broad, flat-bottomed, mature, high-level valley, 
with its floor at nearly 1,000 ft. above sea level, and covered 
with a thick deposit of clay resting on grit and waterworn 
gravel. Remnants of drift material consisting of sand and 
waterworn gravel with occasional large boulders are also found 
at varying heights above the bottom of the valley, and clearly 
do not belong to the present stream conditions. The bound- 
aries of the valley to the east and west are sharply defined by 
two parallel ridges, remarkably straight and of even height— 

the Bull Creek Range, on the east, and the Wickham Hill 


eh 


_ALEXANDRINA 


‘Torrens 
ee 


\ SO a eee eee 


[" waitae! J} Lowlands. 


o 
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ees | Highlands. 


162 


Ridge on the west. The latter is really part of the Willunga 
fault scarp, and its steep western slope is in striking contrast 
to the topography of the valley under discussion. 


The eastern side of the Bull Creek Range also presents _ 


a more broken character, consisting of dissected hill country, 
dropping rapidly to the Murray Plains. These ridges are 
clearly remnants of a once extensive peneplain, dislocated by 
faulting to the west and east. 

They extend northwards beyond the region of the Meadows 
Valley and form the east and west boundaries of the Onka- 
paringa Valley in its upper course. There is no sharply-defined 
watershed between the head of the Meadows Valley and the 
middle course of the Onkaparinga, and the features are such 
as to suggest that an ancient north and south valley of mature 
type and considerable size at one time flowed from the north 
through the Meadows Valley, and the deposits in its floor 
and along its sides also demand such a river. If this be so, 
where was the southern outlet, and when and under what con- 
dition was the present system developed? The upper course 
of the Onkaparinga has every feature of an ancient mature 
valley, and is so regarded by Prof. Walter Howchin. 

To the north of Mount Bold it begins to turn more 
westerly and soon leaves the line of the old valley, which con- 
tinues southerly. The western ridge is breached to the north 
of Mount Bold, and through it the Onkaparinga escapes as 
a deep, steep-sided valley of young character, which it main- 
tains almost to its mouth. | 

Prof. Walter Howchin considers it, in part, an antecedent 
stream whose lower course during the period prior to the 
Mount Lofty uplift was, an ever-changing one over a wide 
fluviatile plain. The extensive deposits of alluvium, gravel, and 
waterworn boulders of the Kangarilla flats and McLaren Vale 
represent, in his opinion, a more southerly course of the river. 
At the time of the uplift its position on the flood plain had 
migrated to that it now occupies, and hence it became incised 
in a rising segment of the highlands. 

The drift deposits of the Meadows Valley, though now 
1,000 to 1,200 ft. above those of the Willunga Plain, strongly 
suggest a dislocated section of that ancient flood plain. 

At that period the ancestor of the present Onkaparinga 
had a more continued north and south direction. The Meadows 
Valley would thus represent a dismembered section of that 
drainage system which, eventually, became choked to the brim 
with fluviatile material. The ever-shifting course of the old 
Onkaparinga tending to a more westerly direction may have 
been assisted by early warping preceding the later block 
faulting. | 


ee 


‘ 163 


There is further a suspicion that there is also a faint 
relic of earlier topography, dating back to the glacial condi- 
tions of Permo-Carboniferous times, now almost obliterated 


x) 
/ Guilt 
£ NC 
. EM, 
== H}| | ; 
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= Wee ae 
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by the later cycles that have been superimposed upon it. 


Brock DIAGRAM 


Highlands showin 


raphy and d rainaye system in relation to 


lopog 


j 
d feclonic control. 


structure an 


Section of southem portion of M! Lofty 


The evidence for this lies in the finding by the writer 
of a glaciated boulder in a pebble deposit in a small road 


164 


cutting, near Dinglebedinga School, in the south-western por- 
tion of the area. This boulder was accepted as glacial by 
Prof. Howchin, into whose charge it was given. Is this a 
fluvio-glacial deposit of Permo-Carboniferous age, or is it a 
Tertiary bed composed, in part, of redistributed glacial? If 
the latter, did it come from the glacial deposits of the Finniss 
River, to the south? This would mean a reversal in direction 
of the present drainage. It is much more probable that the 
glacial deposits of the south, though extensive, are neverthe- 
less a small remnant of a sheet of material which once extended 
much farther north, and the Meadows Valley may, in part, 
be a trough the earliest features of which were due to glacial 
erosion in Permo- Carboniferous times—a much more imperfect 
example of ‘‘fossil glacial topography,’’ however, than that 
of the Finniss River district, described by Prof. Howchin, 
where erosion has laid bare a ‘portion of an ancient landscape 
with remarkable precision. 


What, then, is the past geographical history of this 
region? Briefly it appears to be as follows: —The Meadows 
Valley is regarded as a small dismembered portion of an ancient 
north and south stream the southern course of which, beyond 
the area under consideration, has not been traced but indica- 
tions of it might be expected in the direction of Myponga Creek. 


This valley existed before the uplift of the present high- 
lands, and in the Meadows Creek section there is some pro- 
bability that its course coincided with a much more ancient 
glacial valley partly filled with till. 

Peneplanation advanced to a mature stage with conse- 
quent aggregation and filling up of the old valleys, resulting 
in the formation of an extensive piedmont plain over which 
the streams flowed independently of the underlying structure. 
Early subsidences and warping may have assisted in the insti- 
tution of the diagonal direction of drainage, as shown by the 
present positions of the Torrens and Onkaparinga. Fault 
block dislocation followed, with the gradual establishment of 
the present distribution of highland and plain, giving rise 
to a revived erosion cycle and the entrenching of the deeply- 
cut river valley into the rising segments. The existing cycle 
is one of discordances of level and active erosion along the 
fault scarps, providing short, steep, actively-eroding streams 
tending to cut back into the old topography and divert 
remnants of the old north and south valleys into steep-graded 
easterly or westerly flowing streams. 

The north and south strike ridges of hard rock delayed 
this process, but weak places were eventually found. The most 
important of these was the southern continuation of the Bull 


165 


Creek Range, where the hard Cambrian or Pre-Cambrian rocks 
gave place to softer Permo-Carboniferous deposits. This led 
to an eastern breach by what is now the Finniss River, thereby 
| eapturing the lower course of the Meadows Valley. The 
remarkable course of this river has been referred to by Prof. 
Howchin.“) The same process can be studied on the western 
side, but in a less advanced stage, in several small streams, 
which have actually breached the scarp, but have not yet 
captured much of the drainage of the old valley. The most 
notable of these is Dashwood Gully. Peter Creek, heading 
in the northern Kuitpo Forest Reserve, shows the same 
features. 
BIBLIOGRAPHY. 


(1) 1910—Howcuin, W.: Description of a New and Extensive 
Area of Permo-Carboniferous Glacial Deposits in 
South Australia. Trans. Roy. Soc. S. Austr., vol. 
XXXIV. 

(2) 1913—Howcurn, W.: The Evolution of the Physiographical 
Features of South Australia. Presidential Address. 
Section C, Austr. Assoc. for the Advancement of 
Science, vol. xiv. 

(3) 1918—Tzatz, E. O.: Soil Survey and Forest Physiography 
of Kuitpo. Bulletin No. 6, Dept. of Forestry, 
University of Adelaide. 


166 } 


SOME NEW RECORDS OF FUNGI FOR SOUTH AUSTRALIA. — 
PART II. 
TOGETHER WITH A DESCRIPTION OF A NEW SPECIES 


OF PUCCINIA. 


By T. G. B. Ossporn, D.Sc., Professor of Botany, 
and 

GEOFFREY SAMUEL, B.Sc., Assistant Lecturer and 

Demonstrator in Botany, University of Adelaide. 


[Read August 10, 1922.] 
Puate VII. 


In 1915 one of us published a short note on ‘‘Some New 
Records of Fungi for South Australia.’’ In it were listed 
some forty species the occurrence of which in the State was not 
recorded in the literature dealing with Australian fungi. 

The present paper adds fifty-one species to the fungus 
flora of the State and adds nine names to the host species of 
Australian fungi. Many of these are common, whilst a few 
have already been mentioned in the Annual Reports of one 
of us, and their place in this, list is merely a matter of con- 
venience, since these Reports are difficult of access to most 
mycologists. Others of the species, however, are of more 
interest, because of the apparent rarity of the fungi in other 
parts of Australia, and one of them, Puccima semibarbatae, 
occurring on the native Bulbine semibarbata (Liliaceae), is 
new to science. 

Following the arrangement of the previous list, reference 
is given to McAlpine’s Systematic Arrangement of Aus- 
tralian Fungi, by the number assigned there, and also, where 
possible, to other of McAlpine’s works, in order to render it 
easy to ascertain the range of a species in other States. It 
is hoped to follow this paper shortly by another of a similar 
type, which will bring the published records of South Aus- 
tralian parasitic and micro-fungi into line with local 
knowledge. 

UREDINEAE. 

UROMYCES DANTHONIAE, McAlp. On leaves, leaf-sheaths, 
and panicle-branches of Danthoma semiannularis, R. Br. 
(Danthoma peneilata, F.v. M.,comp. sp.). II., III. Min- 
nipa, Oct., 1916, W. J. Spafford. Also on Danthonia sétacea, 
R. Br., which is a new host plant. South Park Lands, Ade- 
laide, Nov. 2, 1916, T. G. B. O. (McAlp., 1906, p. 85). 


he a 


167 


UROMYCES BULBINIS, Thuem. Teleutosori on flowering 
scapes and leaves, amphigenous, small, densely gregarious, 
frequently concentrically arranged in large circles; at first 
covered by ashen-coloured epidermis, later exposed, firm, 
convex, brown. 

Teleutospores globose to ovate, pedicellate, wall smooth, 


rather thick, 18-25 x 20-224; pedicel deciduous, hyaline to 


yellowish, 3-5 » wide x 3-8 yw long. 
On leaves and scapes of Bulbine bulbosa, Haw. National 
Park, Belair, Sept.-Oct., common (fig. 1), 


Wace vt: 
Teleutospores of Uromyces bulbinis, Thuem. Drawn 
from fresh material (x1100). 


This species was described by Thuemen in Flora, 1877, 
and is given by Cooke, Handbook of Australian Fungi, No. 
1738; also by McAlpine, 1906, p. 87. The localities given 
by the latter are: Victoria, Omeo; New South Wales, Upper 


_ Macquarie River. 


This rust would. appear to be uncommon, for McAlpine 
says he has not seen a specimen. It is abundant on its host 
in parts of the National Park, Belair, affecting the leaves 
and basal parts of the stems, and occasionally sori have been 
found in the flowering region. The appearance is highly 
characteristic, the ashen-coloured young sori showing plainly 
on the yellow-orange infected portion of the host. The com- 
pound sori are occasionally very large, in one case 14x 2 mm.; 
sometimes a second ring of confluent sori surrounds the first. 


168 


We cannot understand the statement in the descriptions _ 


cited above that the sori are ‘‘concave,’’ for when fresh they 


project above the surface of the lesion, whilst the teleiitospores 


in all cases examined by us are distinctly globose or ovate, 
not ‘‘clavate or oblong-clavate,’’ and no case of an acute apex 


has been seen. The spore measurements of the South Aus- | 
tralian specimens are rather smaller than those given by. 


McAlpine and Cooke. Since the fungus is so characteristic in 


its growth, and the only rust affecting Bulbine bulbosa on — 
record, there can be no doubt of the species. Possibly the © 
lack of fresh material by Thuemen may account for the dis- 


crepancies. 
Uromyciapium Trpperianum, (Sacc.) McAlp. On stems 
of Acacia armata, R. Br. Camino in Adelaide district; 


Victor Harbour, Aug., 1915; Athelstone, Aug., 1917, 
Tes GB. es 

On Acacia calamifolia, Sweet. Between Port Augusta 
and Iron Knob, Aug. 22, 1921, J. B. Cleland. Galls 
abundant on the needle-like phyllodes as well as on the pmalles 
twigs. New host plant. 

On Acacia pycnantha, Benth. Millicent, April 7, 1917, 
T. G. B. O.; Meadows, 1921, Ambleside, 1921, y-8: 

The brown potato-like galls of this fungus on trees of the 
golden wattle (A. pycnantha) are so conspicuous that it is 
surprising that no South Australian record of it exists pre- 
vious to 1917. It must have been present some years before 
then, for it was becoming a serious menace to the wattle bark- 
stripping industry in the neighbourhood of Meadows about 
that year. In 1918 severe bush fires swept this area, and the 
wattles which came up after the fire were perfectly free from 
the fungus. During the last year or two, however, galls have 
begun to appear on isolated trees again, and the fungus will, 
no doubt, gradually spread. In this connection it is inter- 
esting to note that trees may be seen loaded with the rust- 
galls, yet surrounded by trees which are perfectly free from 
them. Once a gall has formed on a tree, the fungus spores 
which are produced on its surface probably become splashed 
about in the rain drops, under conditions suitable for germ- 
ination and infection. Thus the tree soon develops numerous 
galls. Spores will usually be carried to other trees, however, 
by the agency of wind, which evidently does not lead to those 
trees becoming rapidly infected (McAlp., 1906, p. 111). 

PUCCINIA BROMINA, Hriks. On living leaves and _ leaf- 
sheaths of Bromus arenarws, Labill. Minnipa, Oct., 1916, 
III., X. McAlpine says mesospores comparatively rare, but 
in this specimen they were fairly numerous (McAlp., 1906, 
p. 116). 


ae 169 


PUCCINIA- FLAVESCENTIS, McAlp. On living leaves of 
Stipa scabra, Lindl., var. auriculata, II. Also on Stipa 
pubescens, R. Br., II., Sept. 23, 1921, W. J. Spafiord. Both 
these are new host species for the fungus. Darluca filum was 
parasitic on the uredosori (McAlp., 1906, p. 119). 


Puccinia semibarbatae, 1. §p. 


Teleutosori on stems and leaves, amphigenous, small, up 
to 2x1 mm., gregarious, occasionally arranged in concentric 
groups, covered by epidermis, convex, becoming exposed, firm 
when fresh, but becoming powdery when dry, deep brown- 
black. 

Teleutospores irregular, fusiform, obovate, or sub-globose ; 
apex generally rounded and not thickened, often conical or 
truncate; rounded at the base, or slightly attenuated; more 
or less constricted at the septum; dark chestnut-brown:; sur- 
face with irregular reticulate ridges and _ depressions; 
33-48 p x 19-26 pu. 

Pedicels short, deciduous, slightly tinted. 

On living stems and leaves of Bulbine semibarbata, Haw. 
Minnipa, Central Eyre Peninsula (S. Austr.), 1915 (fig. 2). 

This rust, which has very characteristic telia, occurred in 
quantity upon the plants of Bulbine semibarbata growing 
around the granite outcrops at Minnipa Hill, on the Govern- 
ment Experimental Farm, Central Eyre Peninsula. It has 
not been found on its host on the eastern side of Spencer Gulf. 

Puccrinia saccaRDolI, Ludw. On living leaves of Goodenia 
amplexans, F. v. M., I., III. Rosetta Head, near Victor 
Harbour, Nov., 1915, T. G. B. O. This host plant is not 
given by McAlpine. The rust is exceedingly common on its 
host in this locality, the aecidia forming large circular patches 
up to as much as 15 mm. diameter on both sides of the leaf. 
The teleutosori occur with the aecidia, usually confluent, often 
forming two, rarely more, concentric rings, usually towards 
the circumference of the aecidial patch (McAlp., 1906, p. 147). 

PUCCINIA ANGUSTIFOLIAE, McAlp. On Podotheca angusti- 
folva, Less., I., I1I., X. Wirrega R.S., Oct., 1916, T. G. B. O. 


By a curious confusion in the synonymy of the host plant, 
McAlpine gives the name as Scorzonera angustifolia, L. The 
error seems to have arisen in the following manner :—The 
genus Podosperma, Labill., 1806, becomes Podotheca, Cass., 
1822, since Podospermum, DC., was already a synonym for 
Scorzonera, L. (Index Kewensis). Podotheca (Podosperma), 
belonging to the Compositae Inuleae-Gnaphalinae, is a genus 
endemic to Australia, and is the host of the native rust 


170. 3 


considered. Scorzonera, of course, belongs to the Compositae 
Chicorieae-Leontodontinae. The native Australian flora con- 
tains but one genus (JMicroseris) belonging to the sub-order 
Chicorieae. 

By the kindness of Mr. C. C. Brittlebank, of the Depart- 
ment of Agriculture, Victoria, we have been allowed to 
examine McAlpine’s type specimens collected at Dimboola, 
Victoria, Nov., 1892. There is no doubt that the fungus 


Fig. 2. s 
Teleutospores of Puccinia semibarbatae, n. sp. (x 1000). 


and host plant from Wirrega (S. Austr.) are the same. 
McAlpine’s note that P. angusttfoliae differs from P. podo- 
sperm, DC., P. scorzonerae, (Schum.) Jacky, and P. 
tragapogi, (Pers.) Corda, in certain particulars is not sur-- 
prising, considering how widely removed the hosts are in 
affinity (McAlp., 1906, p. 150). 

PUCCINIA CALENDULAE, McAlp. On living leaves of 
Calendula officinalis, L., I. Mount Crawford Estate, Jan.,. 
1916 (McAlp., 1906, p. 151). 


‘ 
) 
h 
4 
: 
9 


Se linge seree at 4p 


77 a 


171 


PUCCINIA ERECHTITES, McAlp. On leaves and stems of 
Erechtites quadridentata, DC. Between Coromandel Valley 
and Clarendon, Sept. 23, 1916, T. G. B. O. I. and III. inter- 


‘mixed, mostly on the leaves. A number of three-celled 


teleutospores are present in this specimen. Also Eden Hills, 
Oct., 1917, Miss A. H. Rennie. I., numerous on stems, 
present also on leaves and involucre; III., rare. | 

On Frechtites prenanthoides, DC. Blue Lake, Mount 
Gambier, Oct. 12, 1916, A. G. Edquist. I. only, in groups 
on both surfaces of leaves (McAlp., 1906, p. 157). 

PUCCINIA VITTADINIAE, McAlp. On living leaves of 
Vittadima australis, A. Rich. Wirrega R.S., Oct. 1, 1916, 
T. G. B. O. I., III., and X. on both surfaces of leaves. As 
recorded by McAlpine, the teleutosori were sparsely devel- 
oped; occasionally four or five were observed confluent and 
forming a ring about 2 mm. diameter round aecidia (McAlp., 
1906, p. 164). 

PUCCINIA OPERCULARIAE, (Morr.) Syd. | 

Teleutosori confluent, 3-5 mm. long, forming patches com- 
pletely investing the stem; sori elongate, compact, bullate, 
reddish-brownish, surrounded by the ruptured epidermis. 

Teleutospores golden-brown, oblong to clavate, slightly 
constricted at the septum, smooth, 45-52x15-18y. Upper 
cell rounded, apex thickened, hyaline cap (7-9 ) disappearing 
when germinating. Lower cell about as long as the upper, 
tapering to pedicel; pedicel persistent, 22-50 u long, 3-4 u 
broad. 

Mesospores occasional, similarly coloured to teleutospores, 
fusiform to ovoid, apex thickened with prominent hyaline cap, 
38-45 x 14-18 p. : 

On stems of Opercularia varia, Hook f., III. and X., 
Mount Compass, Oct., 1916, T. G. B. O. (fig. 3). 

This fungus was only found on the stems, where the patches 
of telia formed prominent fusiform swellings, often in the 
middle of the long internodes of the host. The species is « 
Lepto form, the majority of the spores being found with the 
promycelia, or already empty. We have referred the rust to 


‘ P. operculariae, (Morr.) Syd., though it may be that Morrison 


was right in considering this a variety of P. coprosmae, Cooke. 
Groups of confluent teleutosori on the leaves are a feature of 
the latter species; this feature is not noted by McAlpine for 
P. operculariae, nor are mesospores which occur in our speci- 
mens (McAIp., 1906, p. 166). 

PUCCINIA HIBBERTIAE, McAlp. Teleutosori on stems, 
leaves (amphigenous), pedicels, and calyces, causing hyper- 


trophy of stem, densely gregarious, confluent. At first covered 


172 


by the greyish epidermis, bursting, bullate, compact, rounded 
to oval, rarely exceeding 1 mm., chestnut-brown to black. 
Teleutospores elliptical, thickened at the apex, con- 


stricted at the septum, wall smooth, 27-37 x 16-21 p. Pedicel ” 


deciduous, sometimes excentrically displaced, 60-110 p. 

Mesospores occasional, similarly coloured to the teleuto- 
spores, fusiform, generally thickened at the apex, 26-33 x 
12-16 p. 


Fig. 3. A 
Teleutospores and mesospores of Puccinia operculariae, 
(Mgrr.) Syd. Drawn from fresh material (x 690). 


On Hibbertia stricta, R. Br., var. canescens. National 
Park, Belair, Oct. 28, 1916, T. G. B. O. New host species. 

The fungus here described has been recorded under 
McAlpine’s species P. hebbertiae, though it differs from it in 
the size of the spores, which are consistently shorter in our 
specimens. In other respects it conforms to McAlIpine’s 
description (McAlp., 1906, p. 185). 

AECIDIUM OLEARIAE, McAlp. On stems of Olearia 
axtlaris, F. v. M., I., on stems only. Victor Harbour, June 
16,/4918,°T. GG. B. On G@iiecAlp:, 1906; py 197): 


t 


173 


USTILAGINEAE. 


 Ustitago cynopontis, P. Henn. Destroying the inflor- 
escences of Cynodon dactylon, Pers. Mile End, Jan., 1918, 
G. Quinn. 

Although the host plant of this fungus is widely grown 
for lawns, and occurs wild on dunes in many places, the fungus 
had not been recorded in South Australia before. It has 
since been found on many occasions in gardens around Ade- 
laide during the summer (McAlp., 1910, p. 155; Osborn, 
1918, 1921). 

Ustitaco TEPPERI, Ludw. Sori on inflorescences while 
still enclosed in abnormally numerous sheathing leaf-bases, 
forming a compact black mass in which generally only the 
axis of the inflorescence remains of the host. Spore mass 


Fig. 4. 
Spores of Ustilago Teppert, Ludw. (x1100). 


ultimately exposed by decay of the leaf-bases and becoming 
powdery. 

__ Spores brown, globose to ellipsoidal, finely echinulate, 
10-14 pu. 
Bn Neurachne alopecuroides, R. Br. Burnside, Nov., 
1916; Moppa Scrub, Oct. 1917. 

A smut on Neurachne alopecuroides was sent to Ludwig 

‘by J. G. O. Tepper from Torrens Gorge, South Australia, 
who described it in 1889. The fungus appears to have a 
limited Australian distribution, for McAlpine, in 1910, said 
he had not seen any smut on that host. In November, 1916, 
it was first found at Burnside, some six miles south of the 
type locality, and later in the Moppa Scrub, some 30 miles 
to the north. In both localities it is locally common. 


174 


The infected inflorescence is very characteristic (pl. vil., 
fig. 2). The normal host produces an inflorescence at the end 
of a long bare peduncle, as much as 25 cms. above the highest — 
leaf. The lamina of the leaves at the base of the peduncle 
is short (2-3 cms.), with a leaf-base of almost the same length. 
The infected inflorescences have scattered leaves over their 
entire length, and a group of three or more leaves terminating 
the stem. The laminae of these are 1-2 cms. long, with rather 
longer leaf-bases closely investing the diseased inflorescence. 
There is thus a characteristic gall-like development (McAlp., 
1920; ps .161):< 

Cinrracria HYPODYTES, (Schl.) Diet. On stems of Stipa 
flavescens, Labill. Granite Island, Jan. 3, 1919, T. G. B. O. 

Occurring especially on the upper internodes, within the 
sheathing leaf-bases, and preventing the formation of an 
inflorescence (McAlp., 1910, p. 171). 

Urocystis Hypoxipis, Thaxt. On leaf-bases and in- 
florescences of Hypozis pusila, Hook, f. Grange, June 2, 
LOT, TAG BO: 


New host species. The fungus has been recorded from 
Victoria on H. glabella, but it has not been observed on 
the latter in South Australia, though the plant grows com- 
monly in the Adelaide district (McAlp., 1910, p. 197). 


ee ee ee 


? 


es % 


ge ee 


BASIDIOMYCETES. 


AUREOBASIDIUM VITIS, var. ALBUM, Montmart. On leaves, 
young shoots, and inflorescences of Vztis vinifera, L. lLyrup, 
Watervale, -Berri, Renmark, Oct., 1921. ! 

In October, 1921, specimens of young vine leaves were 
sent in from several localities, exhibiting blackened areas of 
irregular extent. If placed in a moist dish, these rapidly 
spread over the whole leaf, and whitish pustules consisting 
of basidia bearing spores on sterigmata formed’ both on leaf 
surface and fruit stalks. In the original description (1882) 
of Aureobasidium vitis, Vial. et Boy., the fungus was described 
as being clear yellow; but in 1897 Montemartini® described 
a variety occurring on leaves and fruit stalks which he named 
A. vitis, var. album, because the pustules were whitish. 
Later in that year, McAlpine described a form occurring in 
Victorian vineyards, chiefly on the berries, as A. vitis, var. 
tuberculatum. The South Australian specimens, both in the 
parts affected and in the nature of the spore pustules, agree 


(1) Montemartini, in: Atti dell’ Istit. botan. dell’ Universita 
di Pavia, 1897 (ref., Zeitschr, f. Pflanzenkrank., vii., p. 359, 
1897). 


init eeinnaiiasiicittintmeeaientinn —% — 
0 OO 


175 


most closely with Montemartini’s description, so that the 
above name is given them (McAlp., 1897, p. 16). 


-ASCOMYCETES. 


ERYSIPHE CICHORACEARUM, DC. On Senecio vulgaris, L.- 
Glencoe, South-East, Dec. 9, 1916, G. Quinn. 

On living leaves and stems of Cucurbita pepo, L. Mur- 
ray Bridge, Feb., 1917. Also common in gardens in Adelaide 
on the marrow and other types of Cucurbita. 


Not listed by McAlpine (Osborn, 1918). 


ASTERINA BAILEYI, Berk. et Br. On living leaves of 
Hakea rostrata, F. v. M. Belair, Sept., 1920, G. S. And 
on Hakea ulicina, R. Br. Forest of Kuitpo, May, 1922, G. 8. 


This is a common fungus, and has been present here for 
years, though not recorded for South Australia’as yet. A 
specimen of H. ulicona, in the Herbarium of the University 
of Adelaide, labelled ‘‘Aldgate, 1895, O. E. Menzel,’’ is 


affected with it. The Hakeas above are new host species 


(McAlp., 1895, No. 1728). 

SEYNESIA BANKSIAE, McAlp. On the upper surface of 
living leaves of Banksia ornata, F. v. M. Forest of Kuitpo, 
May, 1922, G. S. (McAlp., Proc. Linn. Soc. N.S. Wales, 
1903, p. 553). 

PARODIELLA BANKSIAE, Sacc. et Bizz. On leaves of 
Banksia marginata, Cav. Ambleside, May, 1922, G. S. 


This fungus, known as Banksia Freckle, occurs on the 
under surface of the leaves, chiefly the lowest or innermost 
leaves, slightly “languid,’’ as McAlpine says. Although it 
has not been recorded for South Australia before, it has been 
present here for years. A specimen of Banksia marginata, 
in the Herbarium of the University of Adelaide, labelled 
“Aldgate, 1895, O. E: Menzel,” is infected with it (McAlp. 
1895, No. 1741). | 

Oipium, on apple. On living leaves and twigs of Pyrus 
Malus, L. Upper Sturt, Jan., 1921, G. S.; Houghton, Mar... 
Pont, G'S. 

At Houghton the Oidium was unusually plentiful on big 
leafy trees 12 to 15 ft. high which had only been single- 
worked, and were probably on their own roots. 


The perfect stage of this Ordiwm was not found. There 
are four apple mildews—Podosphaera oxycanthae, Podo- 
sphaera leucotricha, Podosphaera tridactyla, and Sphaero- 
theca mali—over which there has been considerable confu- 
sion; until the perithecia are found, therefore, this Oidiwm 


176 


cannot be named. In McAlp., 1895, No. 1721, Podosphaera yf 


tridactyla is recorded for Victoria and New South Wales 
(Osborn, 1919). 


FUNGI IMPERFECTTI. 


SPHAEROPSIDACEAE. 
ConiorHyrium acactaE, McAlp. On living phyllodes of 


Acacia pycnantha, Benth. National Park, Belair, July, 1922, | 


ERS 


DarLvuca FiILUM, Cast. This fungus, which is parasitic — 


on the uredosori of rusts, has not been specifically recorded 


for South Australia before, but has probably often been found. 


Thus McAlpine notes it as common on the uredosori of 
Puccinia lolu, and also gives Mount Gambier, South Australia, 
as one locality for this rust; the same note occurs in his 
descriptions of several other rusts. In our material, Darluca 
filum occurred on Puccinia flavescentis, a host which is not 
recorded in McAlpine, 1906, p. 119, as well as on many 
other species of Puccinia (McAlp., 1895, No. 2087; McAlp., 
HOVG. 0. a2) s 

DrpLop1a ciTRicota, McAlp. Forming scabs on the 
fruit of Citrus aurantium, L. Together with Phoma omni- 
vora, Torrens Park, Mitcham, Nov: 21, 1919. Alone, Clar- 
endon, Jan., 1921, G. S. McAlpine does not record its 
attacking the fruit (McAlp., 1899, p. 83; Osborn, 1919). 

KELLERMANNIA PRUNI, McAlp. Saprophyte on decaying 
almond leaves on the ground, North Adelaide, May, 1921, 
G. S. (McAlp., 1902, p. 104). 

PHoma MAcRopHOMA, McAlp. On twigs of Citrus auran- 
tuum, L. Clarendon, Jan., 1921, G. S. (McAlp., 1899, p. 108). 

PHYLLOSTICTA BRASSICICOLA, McAlp. ‘Ring spot” on 
outer leaves of Brassica oleracea, L. Upper Sturt, Jan., 1921, 
G. 8. (Vict. Dept. Agr. Pamphlet, Cabbage and Cauliflower 
Diseases, 1901). . 


- et 


PYRENOCHAETE ROSELLA, McAlp. Saprophyte on decay- _ 


ing almond leaves on the ground. Blackwood, May, 1921; 
North Adelaide, May, 1921, G.S. (McAlp., 1902, p. 97). 

SEPTORIA DEPRESSA, McAIp. On fruit of Cvtrus auran- 
trum, L., forming circular brownish scabs. Salisbury, Sept., 
1915; Campbelltown, Oct., 1921 (McAlp., 1899, p. 83). 
SEPTORIA DIANTHI, Desm. On living leaves of Dianthus 
carophyllus, L. Fullarton, Sept., 1918, F. W. Eardley. Not 
listed by McAlpine. 

SEPTORIA LEPIDII, Desm. On living leaves of Lepidium 
draba, L. Morphett Vale, Sept., 1915. Not listed by 
McAlpine. 


| 


LT : 


SEPTORIA LYCOPERSICI, Speg. On stems and leaves of 
Lycopersicum esculentum, Mill. Marion, Nov., 1919; Gawler 
River, Dec., 1921. Not listed by McAlpine, but recorded 
for Victoria (C. C. Brittlebank, Journ. Agr. Vict., xvii., 


_p. 498, 1919). 


VERMICULARIA ANGUSTISPORA, McAlp. Saprophyte on 
decaying almond leaves on the ground, North Adelaide, May, 
1921, G. S. (McAlp., 1902, p. 104). 

VERMICULARIA CIRCINANS, Berk. On Alliwm cepa, L. 
Attacking the scales, and spreading occasionally to the green 
leaf portion of seedling onions. Longwood, Oct. 15, 1915. 
Not lsted by McAlpine. 

VERMICULARIA VARIANS, Duc. ‘‘Black Dot’’ disease on 
potato haulms, Mount Gambier, Mar., 1917. On tubers, 
forming slightly sunken areas just under the skin, Carey 
Gully, Jan., 1921, G. S. (McAlp., 1911, p. 92; Osborn, 1921). 


MELANCONIACEAE. 
COLLETOTRICHUM SCHIZANTHI, Jens. and Stew. On stems 


of Schizanthus, sp., causing a wilt. Glen Osmond, Sept., 


1916; Kensington Gardens, July, 1917. Not listed by 
McAlpine. 
GLOEOSPORIUM RIBIS, (Lib.) M. & D. On leaves and 


eanes of Hibes grossularia, L. (Conidial stage of Pseudopeziza 
ribis, Kleb.). Summertown, Jan., 1921, E. Leishman. Not 


listed by McAlpine. 


HYPHOMYCETES. 


ACROSTALAGMUS CINNABARINUS, Corda. Living saprophy- 
tically on decaying potato haulms, forming a reddish mould 
over them. Mount Gambier, April 5, 1917, Ade. i O Nat 
listed by McAlpine. 

CEPHALOTHECIUM ROSEUM, Corda. Developed as a sapro- 
phyte on apple leaves from Ambleside, Feb., 1921, which 
were kept in a moist dish. Commonly develops as a sapro- 
phyte on decaying fruit, and appears to be a facultative 
parasite on stored fruit when the skin is injured. Not listed 
by McAlpine. 

CERCOSPORA APII, Fres. On living leaves of Hees 
sativa, L. (parsnip), causing a leaf spot. Mount Lofty, Sept., 


1919, tT G. B. O. Not listed by McAlpine. 


CLADOSPORIUM PHYLLOPHILUM, McAlp. Dark olivaceous, 


minutely velvety layer over the diseased, wrinkled surface of 
_ peach leaves where injured by Hxoascus deformans. Black- 


i 


wood, Feb., 1921, G. S. (McAlp., 1902, p. 100). 


178 


CoNIOTHECIUM CHOMATOSPORUM, Corda. On twigs of Pyrus 
Malus, L. Blackwood, Nov., 1914, R. Fowler; Mount Gam- © 
bier, July, 1915; Wirrabara, Dec., 1917. Causing cankers on © 
the bark of apples and pears. The severe scabbing of the™ 
fruit by this fungus, which occurs in South Africa and other ~ 
countries, has not been recorded here (Osborn, 1918, 1921). 


CoNnIOTHECIUM scaBRUM, McAlp. On fruit of Cvrtrus 
aurantium, L., causing irregular, flaky, scabbed areas. Ken- 
sington Gardens, July, 1917; Enfield, Mar., 1918; Berri, | 
June, 1922, R. Fowler (McAlp., 1899, p. 80). | 

FumMaGo vaGANS, Pers. On canes of Vitis vinifera, L., — 
forming a black sooty coating, ‘‘fumagine.” Clare, May, © 
19215206. BeOm MeAlp. 91889722238). 


HARPOGRAPHIUM CORYNELIOIDES, Cke. and Mass. Causing 
swollen lesions on the stems of Leptospermum scoparium, 
Forst., with the short, branched, brown conidiophores pro- 
jecting from them. Cleland Gully, near Mount Compass, 
1921, T. G: B, OM(OMeAl py 11895) Nos 1997); 


Orpium oxaLipis, McAlp. On living leaves of Oxzalis 
corniculata, L. Forest of Kuitpo, under ash trees, Dec., 1921, 
G. S. (McAlp., 1895, No. 2276). 


STERIGMATOCYSTIS NIGRA, v. Tiegh. On ripe grapes, the 
skin of which had burst. Southwark, Feb., 1921, G. S. 
(McAlp., 1897, p. 46). 


PHYCOMYCETES. tT 


PLASMOPORA VITICOLA, (B. and C.) B. and deT. “On 
living leaves of Vitis vinifera, L. McLaren Vale, Feb., 1921; 
Watervale, Seven Hills, Berri, and Renmark, April, 1921. 


This fungus appeared first in Australia at Rutherglen, 
Victoria, in the season 1916-17, and in 1917-18 did consider- 
_able damage. From this locality it seems to have spread east- 

ward into New South Wales, and finally Queensland (1920-21). 
Its progress westward of Rutherglen was slow, and not till 
1920-21 did it appear at Mildura. From thence it passed 
down the Murray, appearing at Renmark, Berri, and Water- — 
vale. It was also said to occur at Angaston. The attack | 
was a slight one, evidently resulting from infection late in 
the season. | 

This outbreak is interesting because of the example it — 
gives of the power of dispersal of a fungus disease by wind- — 
borne spores. Mildura, the seat of the nearest epidemic out- — 
break in the past season, lies 100 miles east of Renmark, up | 
the Murray. There is regular traffic between the two places 
by motors, so that it is possible that the spores might have — 


onlay AD Hg 
—— 


el 


179 


been conveyed by human agency or aided by down stream air 
currents along the river. Berri and Renmark are roughly 100 
and 120 miles east of Watervale, and between the places there 
is no direct traffic. Neither is there any direct traffic between! 
the Renmark area and Angaston (in which area downy mildew 


_ is reported) or McLaren Vale, roughly 130 miles south-west 


i 


of Renmark. The chance that spores would be conveyed by 
human agency from the Renmark area to any of these places, 


is very slight. Yet distances of well over 100 miles are con- 


siderable to be bridged by air-borne spores of the Plasmopara 
type. This, of course, is on the assumption that it was from 
the Renmark area that the other South Australian grape- 
growing areas became infected. Unfortunately no reliable 
dates can be obtained of the various outbreaks. They were 
all reported about the same time, except the McLaren Flat 


outbreak in February. It is possible that Mildura was really 


the centre of dispersion for the spores infecting the different 
areas in this State, in which case the carry of the spores would 
be about 200 miles to 230 miles in a straight line. 

No specimens of this fungus have been received during 
the past (1921-22) vine-growing season, although leaves from 
a number of different localities in which the fungus was 
present in 1920-21 have been examined. It seems probable 
that the fungus will have difficulty in establishing itself in 
South Australia because of the climatic conditions (Osborn, 
1921). 

SYNCHYTRIUM TARAXACI, de B. and Wor. On living leaves 
of Hypochoerts glabra, L. Exceedingly common on its host in 
damp areas. Victor Harbour, Aug. 27, 1917; National Park, 
Oct., 1918; Tea Tree Gully, Aug., 1948, T. G. B. O. (McAlp., 
1895, No. 2205). 


BACTERIA. 


PSEUDOMONAS JUGLANDIS, Pierce. On stems, leaves, and 
fruits of Juglans regia, L. This bacterial disease of walnuts 
has, during the last twenty years, spread to almost all places 
in the State where walnuts are grown, even to trees 10 or 12 
miles from any other. It is impossible to get a marketable 
crop from many trees now. Not listed by McAlpine (Osborn, 
1921). 


BacTEeriuM mori, B. and L. Causing angular black spots 


_ on the leaves of Jorus nigra, I.. (mulberry). Clarendon, Jan., 


1921, G. S.; Mylor, Mar., 1922,T.G. B. O. Not listed by 
McAlpine (Osborn, 1921). 


180 


LITERATURE CITED. 
McALPINnE— 


1897—Additions to the Fungi on the Vine in Australia. — 
Vict. Dept. Agr. 
1899—Fungus Diseases of Citrus Trees in Australia. 
Vict. Dept. Agr. . a 
1902—Fungus Diseases of the Stone-fruit Trees in Aus- | 
tralia. Vict. Dept. Agr. 7 
1906—The Rusts of Australia. Vict. Dept. Agr. 
1910—The Smuts of Australia. Vict. Dept. Agr. 
1911—The Potato Diseases of Australia. Vict. Dept. Agr. | 
Osporn, T. G. B.— 
1915—Some New Records of Fungi for South Australia. — 
Trans. Roy. Soc. 8. Austr., vol. xxxix., p. 352, — 
1918—Report of the Consulting Botanist and Vegetable | 
Pathologist, Extract from the Report of the | 
Minister for Agriculture, South Australia, for — 
the year ended June 30, 1918. 
1919—Do., do., for the year ended June 30, 1919. 
1921—Do., do., for the year ended June 30, 1921. 


DESCRIPTION OF PLATE VII. 


Fig. 1. Flowering scapes of Bulbine bulbosa, Haw., showing 
concentric teleutosori of Uromyces bulbinis, Thuem. 


Fig. 2. Two diseased and one normal inflorescence of 


Neurachne alopecuroides, R. Br., showing modifications induced 
by parasitism of Ustilago Tepper, Ludw. 


| 


a 


181 


THE FLORA AND FAUNA OF NUYT’S ARCHIPELAGO AND 
THE INVESTIGATOR GROUP. 


No. 2.—THE MONODELPHIAN MAMMALS. 


By F. Woop Jones, D.8Sc., F.Z.8., 
Professor of Anatomy in the University of Adelaide. 


[Read August 10, 1922.] 


THE FRANKLIN ISLAND Rat. 


The Franklin Island rat was first obtained during a brief 
visit paid to the western island by the s:s. “Conqueror’’ on 
November 23, 1920. The shore party landed shortly before 
noon on a very hot day, and not much life was to be seen 
on the island. An old female and a young male were cap- 
tured a few minutes after landing by clearing out the 
accumulated nesting materials from the hut which has been 
erected upon the northern side of the western island. One 
or two others were seen by various members of the shore 
party, but no more specimens were obtained. ~The two 
animals which had been secured were skinned, but the worst 
possible conditions prevailed for dealing with the material, 
and the skins were by no means good ones. With the cap- 
ture of the first pair a doubt was set at rest, for it was at 
once evident that they were not marsupials, as those who 
knew them best had confidently asserted them to be. But 
though it was simple enough to determine that the animal 
was not a marsupial, it was an altogether different matter 
to establish its identity among the Murines. Its most con- 
spicuous character was that it was a house-builder, and the 
house-building rats were familiar in the literature of explora- 
tion into Central Australia. From the accounts of these 
animals, and especially from an examination of the mounted 
group in the South Australian Museum, it seemed most 
probable that the island rat was Comlurus conditor; and yet 
it obviously differed in some respects from the nest-building 
rat of the interior. It being impossible to proceed further 
with the diagnosis in the absence of type specimens, the old 
female was sent to Mr. Oldfield Thomas, at the British 
Museum. He was good enough to reply at once that the 
animal was not Conilurus conditor, but was a member of the 
genus Leporilus, and possibly was a new species. The second, 
and younger, specimen was therefore sent to the British 


Museum to aid in the establishment of the diagnosis, and 


subsequently the rat was described by Mr. Oldfield Thomas 


182 


(Annals and Magazine of Natural History, ser. 9, vol. viii., 
p. 618, Dec., 1921), and named Leporillus jonesi. When it 
was found that the rat was a new and interesting one it was 
decided to visit the island again, and to arrange for a longer 
stay. The journey was made on the s.s. ‘‘Wookata,” and 
the party camped upon the islands from January 9 to 12, 
1922. Further specimens were obtained, and observations 
and notes were made upon the habits of the animals. Some 
old and bleached skulls were picked up, and plfotographs were 
taken of typical nests. One living specimen was secured, 
but it died as the result of an accident after it had been a 


Figo a: , 
Leporillus jonesi. Characters of the head from a living 
male adult. Natural size. 


week or two in captivity in Adelaide. A third short visit 
was paid to the western island on February 18, 1922, in the 
s.s. ‘‘Conqueror.’”’ On this occasion a few specimens were 
shot, and one was captured alive and uninjured. 


Since the animal has been described by Mr. Oldfield 
Thomas, and will later be dealt with by Mr. E. Troughton, 
of the Australian Museum, to whom specimens obtained on 
the second visit were sent, no attempt will be made here at 
further systematic description. Figs. 1 to 5 depict its most 


183 


important specific characters. Mr. Oldfield Thomas’ account 
is as follows : — 
Leporillus jonesi. 

“Near apicalis, but larger and with shorter ears. Size,. 
as gauged by skull and foot, decidedly larger than in apicalis. 
Fur rather thin and poor, not so thick as in apicalis; hairs of 
back about 17 mm. in length. General colour, above, dull 


es 
a Az 
iy 


1) epee iN 
Y/, 


anovod Tow : 


Fig. 2. 
Leporillus jonesi. Left manus and pes of a female 
specimen. Twice natural size. 


brown (not far from ‘‘Saccardo’s umber’’), the withers tend- 
ing more towards buffy. Under surface slaty-grey broadly 
washed with drabby-whitish, the sides of the belly more 
strongly drabby. LHars shorter than in apicalis, dark brown. 
Hands with the metatarsals dark brown, the digits lighter. 
Feet with the ankles, outer side of the metatarsals (inner 


184 


_ in made-up skin), and proximal parts of digits brown, the 
inner portion of the metatarsals and the tips of the digits 
white. ‘Tail well haired but not tufted, brown above, dull 
whitish below, throughout its length. Not whitened at tip, 
as is also the case with apicalis, the original description not- 
withstanding. 

“Skull larger and stouter than in apicalis. Muzzle broad 
and heavy. Interorbital region broad, with comparatively 


sie 


Fig. 3. 
Leporillus jonesi. The skull from above and below. The 
specimen is from a female. Twice natural size. 


sharp-angled edges. Zygomatic plate more projected for- 
- wards. Palatal foramina short, not reaching the level of m’. 
Bullae rather large—these organs not present in the available 
specimens of agicalis. 

‘‘Incisors rather slender, not thicker than in apicalis, but 
meeting each other at a wider angle, owing to the greater 
breadth of the muzzle. Molars larger than in apicalis, but 
apparently of similar structure; much worn down in the type. 


Ae Fue phen  aainenlagge gaa eas Aho. Moigengyy ten its til dae, og carmella age ap = a ea 


Pe 


ies caer seta as BS hee OY At poet ge We 


185 


‘Dimensions of the type (measured on the skin) :—Head 
and body, 195 mm.; tail, 178 mm. (not quite perfect) ; hind 
foot, 48 mm.; ear (dry), 24 mm. 

“Skull: greatest length, 48; condylo-incisive length, 46; 
zygomatic breadth, 23°5; nasals, 18 x6; interorbital breadth, 
5°7; breadth of brain-case, 18°5; zygomatic plate, 6; palatilar 
length, 13°6; palatal foramina, 88x 3°8; bulla, 7°8; upper 
molar series, 9°3.”’ 

To this description it is only necessary to add one or two 
notes. The fur of the living animal is remarkable for its 
fluffy character, and ‘‘thin and poor,’’ though applicable to 


ki 
UPPER CLEFT 


Fig. 4. 
Leporillus jonesi. Crown form of the molar teeth. From a 
young adult female specimen. Five times natural size. 


the type skin, is not characteristic of the living animal. In 


a state of nature the rat has that compact and fluffy appear- 


ance that is more reminiscent of a little rabbit than of a more 
typical rat. It sits bunched up, so that it appears to be far 
broader and shorter than the prepared skin would suggest. 
The ears are carried well away from the head (see fig. 1), 
and, probably as the result of fighting, they are usually irregu- 
larly notched around their margins. The nipples are four in 
number, and are situated in the inguinal region. It appears 
that the young adhere firmly to the nipples, and for a time 
are dragged about by the mother; it is this circumstance 
which has led to a belief that the animal is a marsupial. 
F 


186 
Measurements of adults, measured in the flesh, are as 


follow :— . 
3 S) 2 g 2 ? 
Rhinarium to eye 21 25 26 26 212° 26 
Rhinarium to ear 40 48 42-5. .* 50 43 5 i 


Ear 7%" ieee 28 24. 30 26 27 
Tail a @ 248. > 162). 162) -.473>,- 4 
Head and body ... 210 240 235 235 198 230 
Hind foot sot ZENS 48 47 44 44 45 
Fore foot Deve i) 19 20 18 19 19 


In the visceral anatomy there are one or two points of 
interest. In the female, the clitoris is completely perforated 


Fig. 5. 
Leporillus jonesi. The palate and 
upper teeth to show the incisive 
papilla and the palate ridges. 


by the urethra; and externally the two sexes are very similar 
in young animals. 

The stomach (see fig. 6) is extremely large, and is very 
distinctly marked out into two chambers by a frilled edge of 


' 

Y 187 

heaped-up epithelium. The first pouch is oesophageal in 
origin, and the second is the true pyloric stomach. The 
caecum (see fig. 7) is enormous; the caput caeci is coiled upon 
itself ; and the whole organ occupies a very large proportion 
of the lower part of the abdominal cavity. In several speci- 
mens it was tenanted by a Cestode which is apparently an 
undescribed species. The small intestine is relatively short, 


Fig. 6. 
Leporillus jonesi. The stomach, showing (a) the outward 
form, and (b) the interior with the well-marked separa- 
tion of the two chambers. Natural size. 


and the large intestine, in addition to its great size, is rela- 
tively long. In Rattus rattus the small gut measures some 
F2 


188 


72 mm., and the large gut some 20 mm.; but in Leporallus 
gonesi the small gut is 57 mm., while the large gut measures 
40 mm. The faceal pellets are more rounded in form than 
are those of the members of the genus Rattus, and they are 
deposited in gropus. 

The rat is a nest-builder, and, so far as I have seen, 
never excavates burrows for itself; in captivity, it shows no 
desire to burrow, or even to scratch into the earth. In the 
islands, a burrow is almost always found beneath the nest, 
and into the burrow the rat will readily retreat; but the 
burrow is always one excavated beneath the nest by a penguin 
(Eudyptula minor) or a mutton bird (Puffinus tenuirostris ). 
There almost seems to be a measure of symbiosis in the economy 


Puls 


SS 


« qaeod Jones 


S29 fee. 
Leporillus jonesi. The caecum. 


A. is the entering small intestine and B. the emerging 
large intestine. Natural size. 


of the rats and the penguins, for practically every nest which 
is found on the northern platform of the islands has a penguin’s 
burrow beneath it. It is a remarkable fact that mutton 
birds, penguins, rats, bandicoots, and the black tiger snakes 
will all bolt into the same hole when alarmed. 

In some of the rats’ nests an enormous amount of material 
is collected, and these large nests appear to lodge a colony. 
Upon the northern side of the eastern island, and high up on 
the cliff, is such a nest; and it is probable that its foundation 
consists of a deserted nest of the sea eagle, the rats having 
invaded it from below. Upon the flat tops of the islands, the 
nests are usually composed of dried herbage, and contain only 


i] 
| 
| 
a 
| 
i 
> F 


189 


a pair of individuals; but upon the island platform they are 
made of sticks of fresh Mesembryanthimum, and nearer to the 
sea of wrack and dried seaweed. The larger nests are com- 
plicated within, a series of passages and chambers being made 
in the heap of collected débris; but the smaller nests consist 
of an entrance run, a central chamber, and an exit run only. 
Upon the sea beaches a whisp of wrack tucked in between 
two boulders, or some seaweed collected in a cleft in the granite 
rocks suffices for a home. In any case, the nests smell badly, 
the lining is stained yellow, and reeks of ammonia; and all 
nests examined were tenanted by a beetle (Hctroma benefica, 
Newm.). 

Quite a large proportion of the rat population lives upon 
the sea beaches, beneath and: between the granite boulders 
which he scattered along the shore. The staple article of 
diet is the succulent leaves of Tetragona implexicoma, and 
enormous quantities are consumed. It would appear that the 
rats also do a certain amount of scavenging along the tide 


Bre. 3. 
Arctocephalus forsteri. External characters of the head 
and face. From a young male specimen. 


line, for their footprints are always to be seen along the 
sand, right down to low-water mark. There is no fresh water 
upon the islands. 

The breeding season is evidently in the colder months 
of the year, for during the time that visits have been paid 
to the islands (November-February) no pregnant or nursing 
females or very young animals have been obtained. 


190 


The rat lives upon both of the Franklin Islands, but upon 
no other islands yet visited. It is by no means nocturnal— 
most of its activities are crepuscular—but at any time of the 
day some individuals may be seen along the shore in the 
intervals between the massive granite boulders. Even in a 
visit at noon, on a particularly hot day, four specimens were 
obtained along a stretch of some 200 yards of beach. There 
appears to be no sort of hostility between the rats and the 
bandicoots (Lsoodon nauticus) which run about and feed 


together, and inhabit the same territory. Indeed, as dusk 


comes on, it is difficult to tell which, among the many shadowy 


forms that appear among the low herbage of the island plat- 


form, is an /soodon and which is a Leporillus. The rats 


are by no means so tame as the bandicoots, and they proved 


to be particularly difficult to take in traps. 


ay 


Pat 3 
hl) 
FRY} Z w \ \ 


ii 
Wi 


Fig. 9. 
Arctocephalus forsteri. . Left forelimb, young male. 


The fur harbours two ectoparasites, a species of flea 
determined by Dr. Ferguson as Hchidnophaga »myrmecobi, 
and a second flea ‘‘apparently indistinguishable from X enop- 
sylla cheopis.” In the intestine cf most specimens is a 
tape worm, which is being investigated by Professor Harvey 
Johnston. . 

Rats of other Islands. 

Goat Island, a waterless island of the St. Francis group, 
is the home of a rat which is evidently abundant; but of 
which no specimen has so far been obtained. - The footprints 
of the rats were to be seen round every boulder upon the 
sea beaches,.and some skeletal remains were recovered from 
the pellets of birds of prey. . It would seem to be a small 
member of the genus Rattus; but all efforts to obtain, or 


a 


S491 


even to see, a specimen failed during the short visit paid to 
_ the island (February 11, 1922). 
a St. Francis Island at one time possessed a-rat, which is 
 gaid to have been quite unlike the house or ship rat, and is 
described as distinctly ‘‘bluish’’ in colour. This species has 
_ long since been exterminated on this inhabited island. 

On Flinders Island is a rat of which.no specimen has so 
far been obtained, but it is almost certainly 2. rattws, since 
it is remembered that the rat-was first seen in the island after 
a vessel had been wrecked upon the shore. 

Pearson Island is probably the home. of two Murines, 
and it is hoped that these species may one day be made 
known to science. ie . 


- Fig. 10 
Arctocephalus forsteri. Left hindlimb, young maie. 


~ 


os oes ae eae ae 


Rabbits. 

Flinders Island alone possesses the unenviable distinction 
of having a rabbit population. These animals were turned 
down many years ago, and for the most part they are black, 
or black and white in colour. It is a great pity that, with 
the continent of Australia as an object lesson, these animals 
should be tolerated on the island, which one day they will 
doubtless overrun. St 


ti ag 
Bed 


pen tees 


Cats. . 


eS 


Cats were liberated many years ago on St. Francis Island. 
For a time they multiplied exceedingly, and have been respon- 
_ sible for the extermination of at least one interesting mar-. 
_ supial species. Of late years they have been decreasing, and 
_ it is to be hoped that the stock is a dying one. 


192 


Seals and Sea-lions. 


There is no doubt the seals that inhabit the islands of 
South Australia are being mercilessly exterminated. A par- 
M\\ 


ia 

i nie wey | i, 
(le 

Wel? wil ne 


Merry 


2 a 


iz 


a 


—— 
Stet ey 


Big: 
Arctocephalus forsteri. Left upper dental 
series of an adult. Natural size. 


tial, but unfortunately a purely nominal, protection is ex- 
tended to them; but the protection goes no further than 


ilies 


193 


the words printed in the Act. Upon the islands where they 
are ‘‘protected’”’ they are slaughtered as freely and as 
barbarously as they are upon the islands where the killing is 
sanctioned by the law. As a matter of fact, the seals anywhere 
upon the islands are at the mercy of any scoundrel who cares 
for the revolting brutality of their slaughter, and deems the 
gain of a few shillings sufficient reward for the labour involved 
in flaying the carcass and preparing the pelt. Only one 
species has so far been seen on the islands. 


Arctocephalus forsteri (Lesson). 
The large ‘‘hair seal’? may still be met with on certain 
of the islands in considerable numbers, and, if no sealing 


party has recently molested them, they exhibit a most 


engaging tameness, evincing a strangely persistent curiosity 
in the coming and going of visitors. There is no need to 
describe the general distinguishing characters of the species, 
and figs. 8, 9, and 10 sufficiently demonstrate the external 
characters of the head and limbs. There are six cheek teeth 
in the upper jaw (see fig. 11) and five in the lower. In colour 
there is a great variety, the variations being apparently due 
to age and sex; but it must also be remembered that, in 
judging the colour of a living seal, the degree of drying of 
the pelage must be taken into account. In the bulls there 
is constantly a lghter-coloured mane. The young pups are 
a rich dark brown, with the naked parts of the skin black; 
the eye is dark brown. 

In the summer months, the seals are for the most part 
in little parties, with pups ranging from some 2 ft. up to 
nearly adult size. The voice of the old bulls is harsh and 
loud, and that of the pups a hissing growl rising to a sequence 
of pathetic yelps very much like those of a small dog. When 
disturbed on the islands most of the animals emit a series of 
long-drawn sniffs, and if the disturbance is continued the 
sniffs become a harsh grunting, and with that the animal 
gallops for the sea. Their pace on land is altogether sur- 
prising, and so is their ability to climb up the steep cliffs 
of some of the islands. On Price Island, especially, are well- 
worn tracks up the cliffs to the top of the island some 250 ft. 
above. On the top of the island, family parties lie basking 
in the sun, and the only danger that a seal is likely to prove 


_is his desire to come down his path to the sea whilst the 


visitor is coming up. Apart from this, they are wholly 
inoffensive animals, and are deserving of all the protection 
that can be afforded them. 


194 


FLORA AND FAUNA OF NUYT’S ARCHIPELAGO. 
No. 3.—A SKETCH OF THE ECOLOGY OF FRANKLIN ISLANDS 


By T.-G. B. Ossorn,: D.Sc., 
Professor of Botany in the University of Adelaide. 


[Read August 10, 1922. ] 


Puates VIII. to XI. 


The following sketch of the ecology of Franklin Islands 
embodies the results of a brief visit paid to the group in 
January of this year. It had been the intention of the party 
to spend over four days ashore, but owing to adverse weather 
delaying the ship the time was reduced to two and a half; 
this shortage of time, and the season of the year, — explain 
any obvious imperfections in the account. 


GENERAL. 


Franklin Islands “) form a small group consisting of two 
main islands with some outlying rocks and islets lying in 
lat. 32° 27' S., long. 133° 39’ E., some 12 statute miles off 
the nearest mainland. The largest islands are each about 
one and a half miles in length, are flat-topped, and joined 
together by a sand-bar which dries at low tide. The western 
island is 159 ft. high and the eastern nearly the same height 
A chain of rocks about one and a quarter miles in length, 
some of which are above water and the highest elevated about 
15 ft., les about half a mile off, and nearly parallel to the 
south coast of the western island. A pyramidal islet about 
50 ft. high standing on a rock platform which dries at low 
tide extending nearly 400 yards from it, lies 1 »200 yards east- 
ward of the eastern island (pl. vii, fig. 1); 

Though not, strictly speaking, a part of Nuyt’s Archi- 
pelago, the Franklin Islands were sighted from it by Matthew 
Flinders in 1802 and were named by him. Flinders and his 
party did not, however, land upon them. Had they done so 
it is probable that Robert Brown would only have received 
further confirmation of his opinion as to the sterility of the 
islands along the central part of the south coast of Australia. @ 


(1) Australia Directory, 10th edit., vol. i., p. 149, 1907. 


(2) Brown, R., Botany of Terra Australis. Appendix to 
Flinders’ Voyage, vol. ii., p. 534. Point Brown, one of the nearest 
portions of the mainland to the east of Franklin Islands, was so 
named by Flinders in February, 1802, in honour of Robert Brown, 
naturalist on board the ‘Iny estigator.’ 


és 


PS, 
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195 


The Franklin Islands form a part of the pastoral lease 
of Mr. Lloyd, to whom [ am indebted for information as 
to his impressions of their climate, etc. They are uninhabited, 
there being no fresh water upon them, but have been used 
in the past as grazing for a few sheep. Since the 1914-15 
drought they have not been grazed, and at present there is 
little sign of disturbance owing to human occupation, even in 
and about the small stockyard erected near the anchorage. 
The islands can seldom, if ever, have been visited by a botanist 
before, and in their present condition it may be fairly assumed 
that they present a reasonably complete exhibition of their 
original vegetation. The influence of the white man is seen 


am the presence of a few alien annuals, but in January these . 


were not much in evidence. 


PHYSIOGRAPHIC FEATURES. 

‘No account of the general geology and physiography of 
the Nuyt’s-group has been published. Howchin,® however, 
has visited the islands eastward of Cape Catastrophe, and 
from his account it would-seem that Franklin Islands are 
essentially similar. The islands described by Howchin rest 
on a platform of remote age (Cambrian or Pre-Cambrian) 
formed of an intricate series of metamorphic, volcanic, and 
plutonic rocks of deep-seated origin. These old rocks le a 
few feet above or below sea level and represent a base level 
of erosion, considered marine. On these platforms is a cap- 
ping of very recent date (post-miocene) which consists of 
wind-blown sand, formed at a time when the sea was retreating 
south of the present coastline. This sand has become in- 
durated owing to the action of rain water on its calcareous 
contents. “‘In times immediately antecedent to the present 
the sea returned to its old areas, and is now washing away the 
soft wind-constructed sandstones that were left in the line 
of its former retreat.’’ 

The solution of calcareous matter in the soil and its sub- 
sequent deposition, as the water evaporates, in the form of a 
bed of travertine limestone below the surface is a marked 
feature of such areas. 

The Franklin Islands, so far as it was possible to observe 
them, agree with the type of geological formation described 
above. The platform of the islands is granitic, on which rests 
more or less consolidated sandstone. In one or two places 
immediately above the granitic platform a thin deposit of 
pebbles suggests the occurrence of a conglomerate. The cliffs 
at the north-west end of Eastern Franklin are decidedly more 


(3) Howchin, W., Proc. Roy. Geogr: Soc. S. Austr., x., pp. 
‘204-219, 1909. 


196 


clayey than in other places, but nowhere was any superficial 
deposit of clay noticed as forming a compacter and more 
retentive soil. The surface soil is generally white sand, which 
is almost everywhere exposed owing to the open vegetation. 
The soil types may be grouped as more or less consolidated 
sand, travertine limestone, and white drifting sand. 

The foreshore is of two types, rocky or sand. The whole 
of the way along the south and west coasts, and along much 
of the east, too, the waves wash over a broad pavement of 
granitic rock that slopes away at a low angle to the sea, or 
they beat upon a jumbled mass of boulders caused by its 
destruction. There is thus no room for the development of 
a littoral flora along most of these coasts, for the consolidated 
sands rise from the platform of rock at a steep angle to the 
plateau summit. Only in a few places does the development of 
a wider boulder breastwork allow of the accumulation of a 
little sand at the cliff foot upon which littoral plants appear. 
On the south coast of the Western Island the cliffs rise 20 to 
30 yards back from the beach, the intervening area being a 
level stretch of sand raised some 6 ft. above the shore line. 
The terrace thus formed is a curious and distinctive feature 
of the island, that suggests at first sight a raised beach, but 
which is capable of other explanation. 

True dunes are developed only at the north-east end of 
each island, especially the western, near the sand-spit that 
connects the two islands. The strong south and south-west 
gales sweeping round the corner of the islands deposit the 
sand in these comparatively calm areas, building up a small 
but typical coastal dune of the unstable type. 

The summit of the islands is a gently undulating plateau 
termed’ the ‘‘roof’? in this sketch. The southern coast is 
highest, from whence there is a slight slope downwards towards 
the north, the highest point on the group (159 ft.) being a 
rounded knoll lying near the south-west corner. » 

The partly consolidated sands of the roof and cliffs of the 
islands are honeycombed*:by burrows of mutton birds or sooty 
petrels and penguins. The effect of their activities is to con- 
stantly disturb the sand between the larger bushes and open 
the way for wind erosion. The sand which is blown away is 
either held by vegetation on the roof forming local white 
dunes, or is blown to the lee-side of the island and washed 
away. ‘The terrace referred to above on Western Franklin 
has possibly been formed by such an accumulation of wind- 
blown sand. 

Once wind erosion starts at any point the effect is cumu- 
lative, and a ‘‘blow out’’ may develop, as it would in a recent 
sand-dune area. Several such areas can be seen in various 


£07 


stages upon the roof, especially upon Eastern Franklin 
(pl. viii., fig. 2). When the superficial layers are removed the 
underlying travertine is exposed as a pavement, which resists 
erosion. Local patches of travertine that have been exposed 
in this way may remain as knolls rising a few feet above the 
general level of the roof (pl. ix., fig. 1). Travertine pavements 
bear a characteristic flora that by its growth leads to their 
disintegration, when the sand flora reappears. 

Nowhere on the islands is there anything in the nature 
of a watercourse, claypan, or rockhole. All rain that falle 
must sink directly into the soil, and presumably soaks through 
to the granitic platform. It might be expected that along 
the edge of this there would be damper areas, or even springs, 
but if this be so they were dry in January. 


i 
{ 
| 
| 
| 


CLIMATE. 
; = Meteorological data of uninhabited islands are obviously 
| difficult to obtain, but some impression of the climate can be 
gained by comparison with the mainland nearby. The two 
nearest stations of the Commonwealth Meteorological Service 
| are Fowler Bay and Streaky Bay; the records of these are 
| given below by the courtesy of the State Meteorologist, to 
whom my thanks are due. 
| Tasie I. -! 


! Average, highest and-lowest monthly rainfall at Streaky 
Bay (S.) and Fowler Bay (F.), in inches, for a period of 44 


years :— 

te = Sy Oo ~ . yas ‘ S . 5 

a ra a u 3 fs} we oO aA |r z 3) = 

alslalelej2l2l2 le lg lz lz] 2 

| | 
Rirerd ew IS: 66.54: .43| .54] .59]1.02]1.97|2.86|2.36]1.94|1.36] .95] .68] .40/15.10 
= sao .38| .50| .50] .87/1.82/2.19]1.74|1.47| .94| .87| .60| .30]12.16 
Mighiest, § 6.2003. 3.37|4.67|2.43|4.05|4.81|7.51/6.02|5.12|4.03|2.37|4.18|2.48|23.50 
Bis cccgest.v: 2 4(5.90|3.26)3 7431/5768. 305.8212. 62)2.6712.79)143l19.0 
HBOS Son slec case 00] .00} .00| .00] .13] .26] .56] .43] .13| .00| .00| .00} 9.34 
WE a .00} .00| .00} .00] .31 ae .32 ie oe oo .00| .00] 6.91 
TaBLe IT. 


Average, absolute highest and absolute lowest temperatures, 
in degrees Fahrenheit, at Streaky Bay (S.) and Fowler Bay (F.), 
The records at Streaky Bay have been kept for 31 years, and for 
six years at Fowler Bay :— 


3 5 = by . . . ww 

ee ae ea ee begets 

o Co) = 3) cS) cD) 
ee OU Shi mend | alte ine. (ah) BO zai Gasllapsl, 
Ay. temp. S.| 71.7] 72. é 68.5] 63.9 saalsa.s}sz.9 54.9/57.8] 62.5| 67.0 7a. 67.2 
68.9| 69.2] 68.0| 63.6|59.9/54.2/53.6|55.4|58.7| 61.6| 64.7] 68.1] 62.2 
Abs. high, S.]114.2/114.2]104.7] 96.0]88.3]79.0|73.0]83.0|91.0/104.2|113.8]117.0]117.0 
F.|109.0]113.0/108.5]100.2/91.3/82.8|79.5|/86.0/95.0/108.8/114.5/113.9]114.5 
Abs. low, S.| 46.2] 44.8] 43.5] 40.2/34.0/32.0/31.2|32.2/33.9| 38.0] 39.5| 42.5| 31.2 
««  F.} 50.5} 50.5] 49.5] 42.0/36.1|35.0/32.1/35.1|36.5| 40.0] 47.0| 48.7] 32.1 


ae, | “4 


: 
| 
| 
| 
| 
| 
| 
| 


198 


From the foregoing tables it is seen that the climate is of 
the semi-arid type,(4) though as the Franklins are islands the 
conditions are naturally less severe than on the mainland. Mr. 
Lloyd, owner of the lease, says that in his 40 years’ experience 
on St. Francis Island (one of the Nuyt’s Archipelago group) 
he has not known either a frost or a day of over 100° in the 
shade. The atmospheric humidity, also, will be greater on 
the islands than on the mainland. There was ample evidence 
that wind-shearing had an important effect on the growth form 
of the plants, and in killing back the exposed shoots of such 
relatively xerophytic plants as Olearia axillaris and Calo- 
cephalus Browmi. 

As regards the rainfall (Table I.), it will be seen that 
the records of both mainland stations show that no rain may 
fall during seven months out of the year, and that the average 
precipitation for the four months December-March is ‘50 in. 
or less per month. : 

VEGETATION. 


Inttoral Flora.—This may be considered under the head- 
ing of fore-cliff vegetation and coastal dunes, but except on 
the north coast is everywhere sparsely distributed. The main 
fore-cliff vegetation is Calocephalus Brownw, the individuals 
of which form low rounded bushes, | ft. to 25 5 ft. high. The 
branching is densely intricate, and the linear-cylindrical leaves, 
3 mm. long, stand erect, parallel to the stems. The whole 
plant is white in colour owing to the development of close 
tomentose woolly hairs. In places, along the north coast 
especially, and less frequently along the other coasts, such 
. bushes form a continuous line a yard or two wide, fringing 
the shore, rooting in the coarse sand, and often protected on 
the seaward side by granite boulders. Occasionally bushes 
of Myoporum insulare and Nitraria Schoeberi are to be found 
in association with the Calocephalus, when the strip of vege- 
tation is wider. Both the Calocephalus and Myoporum showed 
obvious wind-shearing, the twigs on the weather side being 
cut back and dead. 

Other developments of a Tittoesl flora, except in the case 
of dunes, can be considered as being rather in the nature of 
accidents than characteristics of the habitat. As such may 
be considered the occasional patches of Frankenia pauciflora 
developing at the foot of the cliffs in the sand that had 
lodged behind a wide zone of granite boulders. More character- 
istic of the littoral habitat was the development of a pure 
sward of Sporobolus virgimcus just above the high-tide mark 


(4)Cannon, W. A., Plant Habits and Habitats in the ea 
Portions of S. ‘Austr., Carnegie Inst. Pub., No. 308, p. 2, 1921. 


iat 
We 


e 
. 
tf 


199 


along one or two long clefts (dykes of softer rock) in the 
granite. This was the only habitat of the plant observed upon 
the island. 

Coastal Dunes.—The dune formation is of the typical 
South Australian type,®) but is poorly represented, and is 
best seen in an active state at the north-eastern ends of the 
islands, as has been noted above. The first colonist is, as is 
usual, Spinifex hirsutus, which grows (though not abundantly) 
at the east end of Western Franklin. The strong wind action 
was shown by long tails of drift sand behind the Spinifex 
clumps. The usual succession towards dune shrubland is 
shown, Syinifex being followed by Olearia, though no thickets 


develop. Together with the Olearia there are bushes of 


Myoporum imsulare, these especially showing wind-shearing. 
The dune flora exhibited is of a depauperate and shifting dune 


type. Scaevola.crassifolia, found elsewhere on the island, does 


not enter into it, as is commonly the case on the mainland, 
while Leucopogon Richer and Muehlenbeckia adpressa—the 
latter very common on the mainland in the immediate vicinity 
—were not observed. An atypical dune formation is developed 
fairly commonly along the north coast, and merges into the fore- 
cliff flora described above. Along this coast there is no fore-dune 
flora, for the sand does not blow up from the sea, but is carried 


across the island by the south or south-west winds, and is 


deposited at the foot of the slopes. In one case a terrace 
some 20 yards across, and raised 4 to 6 ft. above the general 
level of the shore, has been formed. The coastal face of this 
falls steeply to the shore, and is a surface of erosion rather 
than apposition. This factor may determine the infrequency 
of Spinifex hirsutus, only-found in one small patch, whereas 
on the normal dune it is the pioneer plant. Shrubs of the 
sand-dune type are represented by occasional bushes of Olearia 
axillaris, whilst, where there are more granite boulders and 
less sand, Myoporum insulare, Nitraria, Calocephalus, and 
Scaevola also come in. The greater part of this terrace is 
covered more or less completely by Mesembryanthemum 
aequilaterale, Threlkeldia diffusa, and Enchylaena tomentosa, 
which form an open association much disturbed by the burrows 
of mutton birds and the tracks of penguins. At one end of 
this area a small hut and shearing shed with sheep yard fenced 
by posts and wire has been erected. The amount of disturb- 
ance of the vegetation caused by this is exceedingly slight. 
The most noticeable feature is the way that Tetragoma an“ 
Enchylaena grow as scrambling’ climbers over the posts an: 
wires, so that the fence resembles a low hedge in places. 


(5) Osborn, T. G. B., Brit. Assn. Rep., Australia, 1914, pp. 584-6. 


200 


Cliff Vegetation.—The word ‘‘clifi’’ is here used to in- 
clude the various types of slope rising from the sandy beach 
- and granite platform of the island to the roof. In most places 
these slopes are not steep enough to be termed cliff in the 
ordinary sense of the word, especially where they are sandy; 
but in others, where they are formed of a denser clayey 
material, the slope is too steep to be climbed easily. It is 
convenient to use the term cliff, with the reservation as above, 
when speaking of those sandy slopes from the roof to the 
shore, which are composed of consolidated sands, to dis- 
tinguish them from the blown sand on which the littoral 
flora is developed. 

It was not possible to distinguish any special associations 
on these areas. The flora they bear is essentially the same as 
that of the roof adjoining, but growing under more exposed 
conditions. The most frequent type of vegetation is a com- 
munity with Mesembryanthemum australe dominant (pl. x., 
fig. 2). This is often a monospecific community, especially on 
the exposed faces of the south coast. Mesembryanthemum 
australe grows commonly on the mainland at the margins of 
salt swamps (in distinction to Mesembryanthemum aequila- 
terale, which is psammophilous), and its dominance on these 
exposed cliffs suggests that they are often wetted by spray 
during high winds. On faces less exposed to spray Mesem- 
bryanthemum australe grows with Frankema pauciflora. The 
impression was gained that the relative proportions of these 
plants offered some rough idea of the degree of exposure and 
consequent salinity of the soil at the spot. 

Salsola kali was found to be dominant, and often the 
only plant, on some cliff slopes where the sand was less con- 
solidated. The cliffs of the north coast, especially on Eastern 
Franklin, show greater diversity of flora. It is probable that 
here, the exposure to spray and wind being less, the soil factor, 
with its consequent effect on drainage and aeration of roots, 
has more play. The communities observed were all open, but 
it was possible to recognize more than one type. On steep 
slopes, where the sand was mixed with some amount of clay, 
Nitraria, Mesembryanthemum australe and Frankema fruti- 
culosa were most abundant, Threlkeldia, Hnchylaena, and 
Mesembryanthemum aequilaterale also being present. At one 
point a small landslide had taken place recently, and Mesem- 
bryanthemum australe and Frankenia fruticulosa were noticed 
as first colonists of the newly-disturbed ground. 

Where the cliff face was more sandy (pl. x., fig. 1), 
Scaevola, Myoporum, and Olearia develop with Mesembryan- 
themum australe, Threlkeldia, and Frankema fruticulosa as 
ground plants. It is probable that in such places Scaevola 


pee 


201 


_ obtains conditions nearest to those under which it grows on 
the mainland. Locally it may almost be said to become 
dominant, but it quickly disappears where the soil is more 
compact. 

Vegetation of the Roof.—There are three main associa- 
tions to be recognized on the roof of the island, but in the 
limited time that was available to examine them it is difficult 
to describe them in other than a static sense. Their possible 
‘relation to each other as members of a succession will be 
discussed shortly below. 

(2.) Rhagodia crassifolia, open shrubland.—This is the 
most stable type of association seen upon the island. Low 
bushes of Rhagodia crassifolia, 1 ft. to 2 ft. high, cover con- 
siderable areas, this being almost the only species in the com- 

_ munity(pl. xi., fig. 1). The association is an open one, with bare 

patches between the bushes, but it is thought that biological 

_ factors in the form of mutton birds and rats are largely respon- 

| sible for this, and that, if these were removed, the covering 

of Khagodia crassifolia would quickly be complete. The white 
ground seen in the foreground (fig. 1) is caused by mutton 
bird burrows, while in other places the Franklin Island rat 

_ (Leporillus jonesi, Thos.) had gnawed down portions of bushes 

near to the ground and used the stems to construct its house 

| or wurlie, around other living bushes (see No. 2 of this 
| 


series.) 

The only other plant noted in this association was 
Stlozerus tomentosus, a small deep-rooting annual, the 
numerous wiry stems of which grow at first prostrate, then 

turn erect, and are terminated by characteristic compound 

_ capitula of yellow flowers. 

The prevalent colour of the vegetation here in January 
was a dull grey-green. 

(u.) Frankema fruticulosa association on travertine pave- 
ments.—Pavements of travertine or nodular limestone occur 
at all levels from a few feet above the shore line to high points 
on the roof. They all bear a characteristic flora of which 
Frankema fruticulosa is the most typical species (pl. ix., fig, 
1; pl. x., fig. 2). This is a mat-forming woody plant, which, 
though it sometimes forms a small tap root, also develops 
adventitious roots freely on the underside of its prostrate 
stems. The stems, except in the oldest parts, are hidden by 
numerous opposite linear-cylindric leaves (3 to 4 mm. long) 
standing erect. The leaves are almost grey owing to hairs, 
with only a tint of subdued green. They are also revolute, 
showing a pronounced groove on the under-surface. The thin 

+ wiry roots run horizontally at no great depth in the sandy 
soil between the limestone blocks. They have a solvent action 


202 


on the limestone, so that the upper-surface becomes etched by 
their growth and finally eaten through. 

Two annual grasses grow in the sand-filled cracks of the 
limestone pavements. They are Danthonia setacea and 
Calamagrostis filiformis, of which the -former is most 
abundant. At low levels near the sea Mesembryanthemum 
australe and Tetragonia implexicoma also occur; the latter was 
only seen along cliff edges, over which it scrambled. 

Where the pavements are covered by sand, WNtraria 
Schoebert develops a more or less extensive mound owing to 
its growth habit (pl. ix., fig. 1). With it Olearva axillaris, 
and occasionally Stipa teretifolia, become associated. These 
plants represent a colonization of the pavement by the flora 
of unstable sand, owing to the Witraria; they are not typical 
of the flora of the limestone pavement as such. 

Frankema fruticulosa is certainly the character plant of 
this association. It gives a most characteristic appearance to 
the areas, which can be distinguished from a distance by their 
light-grey colour (pl. x1., fig. 1). The stability of the associa- 
tion is limited by the existence of the limestone. As this is 
broken up by the solvent action of the roots or rain water, 
the proportion of sand exposed becomes greater, leading to 
an increase in the number of annuals, and also such sand- 
collecting bushes as Witraria and Olearna. Ultimately, there- 
fore, the travertine pavement flora is replaced by an open 
shrubland passing through a phase in which the proportion 
of grasses 1s greatly increased. Such a transition was noticed 
on the roof of Western Franklin in a ridge of sand with 
limestone rubble, bearing old plants of Frankema fruticulosa 
and much Danthonia pemcillata, which is a perennial. 

(iii.) Open association on loose sand.—At present about 
half the roof area is occupied by an open association in which 
the most prominent plants are Salsola kali, Lepidium foliosum, 
and tussocks of Stipa teretifolia, the only perennial plant con- 
stantly present (pl. ix., fig. 2). With these also occur Bromus 
' arenarius, Silorerus tomentosus, Vuittadima australis, etc. 
Such areas are literally honeycombed by the burrows of mutton 
birds or penguins, so much so that they are unpleasant to 
walk across, as the ground constantly caves in under-foot. The 
soil is, therefore, constantly disturbed, and large areas are 
bare, though annuals probably occur in the winter months. 
As our visit was paid in January no list of this therophyte 
flora could be made; from the fruits collected under bushes 
it was clear that an introduced Hordeum occurred, and also 
Daucus brachiatus. Nitraria Schoebert plays a prominent part 


in this association in some places (pl. viii., fig. 4) 5 it develops : § 


(6) Cannon, W. A., be., p. 70. - 


203 


dunes about itself, which, as the original bush dies away, 
become colonized by. Olearia aaillaris and Frankenia 
paucifiora. F 

Several patches; some up to half an acre in extent, were 
strewn with the dead stems of Lavatera plebeja.- Professor 
Wood Jones says he saw thickets of this plant when he landed 
on the island in December, 1920. One place indicated by 
him as a locality in which Lavatera was specially dense is now 
an area of shifting sand (pl. xi., fig. 2). It is believed that 
the growth of the mallow was largely responsible for this. On 
sand the plant is mainly biennial, and by its dense growth 
would tend to kill out the ground vegetation below it. When 
the Lavatera dies there is nothing left to hold the soil, and, 
as a result, the sand drifts. 


SUMMARY. 


Plant Succession.—The general trend of succession can 
only be briefly suggested. The Rhagodia crassifolia shrub- 
land is the most stable community, and probably represents 
the climax on the island. It is not, unlikely that this associa- 
tion is really a subclimax, climatic factors limiting the suc- 
cession, which, to judge from the mainland nearby, one would 
expect to reach a scrub woodland composed of mallees 
(Eucalyptus spp.) and Melaleuca parviflora. Though the 
Rhagodia crassifolia shrubland is regarded as a climax, no sign 
was seen of its spread or regeneration, rather the reverse. 
Biotic factors, notably the burrowing of birds, operating with 
the wind factor, disturb the area and tend to the development 


of drifting sand. Ultimately, if the present set of factors 


remain unchanged, the whole of the consolidated sands will be 
removed and the bare granite platform be left. The last 
stage of the Franklin Islands will be a wave-swept reef similar. 
to that lying immediately south. This is clearly shown by the 
intermediate phase illustrated on the eastern islet before- 
mentioned (pl. vili., fig. 1). 

If there be little sign of. regeneration of the climax there 
are earlier stages in succession to be seen. The clearest is the 
passage from the Frankenia fruticulosa association through 
a mixed low shrub and grassland with Wrankenia pauciflora 
and Danthoma penicillata to a mixed open shrubland of 
Nitraria, Olearia axillaris, Enchylaena, Threlkeldia, etc., with 
Stipa teretifolia and various annuals. This association is un- 
stable. The action of birds and wind depresses the succession 
to the Stipa, Salsola, Leyidium community described. This 
unstable association is apparently gradually coming to occupy 
most of the island. The most important sand stabilizer at pre- 


sent is Vitraria Schoeberi, which, owing to its dune-forming 


204 


capacity and. high salt toleration, tends to maintain a shrub- 
land as opposed to an open community of tussock grass and 
annuals. 

Flora.—Considering the interest of the fauna, it is rather © 
remarkable that the flora should be so limited and without 
any peculiar species. A complete list of the flowering plants 
collected is given below. It numbers only 34 species, though 
owing to the season the annuals are probably incomplete. The 
list includes eight grasses, six composites, and five Cheno- 
podiaceae. The complete absence of Leguminosae and 
Myrtaceae is surprising. The neighbouring coast has Acacia 
spp. on the dunes and mallees (Hucalyptus spp.) and Mela- 
leuca parviflora on the consolidated sands. Itissaid that these — 
plants occur on some of the neighbouring islands, which are 
larger. The present flora of the Franklins is probably vestigial, 
but there is no evidence that it included more Phanaerophytes 
in recent. times. 

Considering the flora in regard to the growth-forms, it 
will be noticed that there are five species of shrubs (14%), 
13 species of undershrubs or perennial herbaceous plants 
(38%)—all chamaephytes—16 species of annuals (45%). Dis- 
regarding the percentages, which are probably misleading 
owing to the very small total number of species, and incom- 
pleteness of the annual (therophyte) flora, it will be seen that 
there are no Phanaerophytes other than Nanophanaerophytes, 
and that the whole classes of Hemicryptophytes and Crypto- 
phytes are absent. This indicates the severity of the environ- 
mental factors, especially wind, as regards the absence of the 
first, and edaphic conditions (unstable soil) as regards the last 
two groups. The aridity of the environment is indicated by 
the relatively large number of annuals (Therophytes), which 
is almost certainly understated in the list. | 


s 


APPENDIX. 
The following is a complete list of the plants observed or 
collected. I am indebted to Mr. J. M. Black, who has kindly 
assisted in determining some of the species :— 
N.=Nanophanaerophyte ; Ch. = Chamaephyte ; 
Th. = Therophyte. 
Spimfex horsutus, Labill. Ch. 
Stipa teretifolia, Steud. Ch. 
Sporobolus virguucus, Kunth. Ch. 
Calamagrostis filiformis, (Forst) Pilger. Th. 
Danthonia pemcillata, (Labill.) F. v. M. Ch. 
Danthoma setacea, R. Br. Th. 
Bromus arenarius, Labill. Th. 


—_—e. , 
> ~ 
t ; 
. . A 
or. . 
4 Ramey . 
= 


205 


*Hordeum sp. (seed only). Th. 

Dianella revoluta, R. Br. Ch. 

Rhagodia crassifolia, R. Br. N. 

Atriplex prostratum, R. Br. Th. 

Enchylaena tomentosa, R. Br. Ch. 

Threlkeldia diffusa, R. Br. Ch. 

Salsola kalt, L. Th. 

Mesembryanthemum aequilaterale, Haw. Ch. 
‘Mesembryanthemum australe, Sol. Ch. 
Tetragoma impleaicoma, Hook, J. Ch. 
Leyidium foliosum, Desv. Th. 
*Sisymbrium orientale, L. Th. 

Nitraria Schoebert, L. N. 

Lavatera plebeja, v. tomentosa, Sims. Th. 
Frankema fruticulosa, D.C. Ch. 

Frankema paucifiora, D.C. Ch. 

Daucus brachiatus, Sieb. (from fruits only). Th. 
Nicotiana suaveolens, Lehm. Th. 

Myoporum insulare, R. Br. N. 

Scaevola crassifolia, Labill. Ch. 

Olearia axillaris, F.v. M. N. 

Vittadima australis, A. Rich. Th. 

Siloxerus tomentosus, Wend. ‘Th. 
Calocephalus Brownu, F.v .M. N. 
Gnathalium luteo-album, L. Th. 

Senecio lawtus, Sol. L. (a hairy form as well as the usual 

glabrous one). Th. 
*Sonchus asper, All., v. httoralis, J. M. B. Th. 


DESCRIPTION OF PLATES. 

Puate VIII. 

Fig. 1. Islet off Eastern Franklin, showing the granitic 
platform with a small cap of consolidated sandstone at one end. 
The slope in the foreground is a travertine limestone pavement in 
process of decay. Frankenia fruticulosa, accompanied with Dan- 
thonia setacea, is being invaded by Stipa teretifolia, Frankenia 
paucifiora, Threlkeldia diffusa, etc. Large patches of Mesembryan- 
themum australe present. 


Fig.2. Area on south coast of Eastern Franklin, showing sand 
drifting away from and exposing travertine pavement. The blown 
sand is held by Nitraria Schoebert. The mounds in the foreground 
are formed by Frankenia pauciflora, which holds the sand forming 
mats up to 2 ft. in diameter. Beyond Rhagodia crassifolia shrub- 
land with occasional travertine pavements. In the middle distance 
is another “blow-out’’? area with Nitraria, Frankenia pauciflora, 
and Salsola kali. 


An * denotes a plant not indigenous to Australia. 


= 
o~ 


she 


206 


weer 


PLATE IX. 


Fig. 1. Travertine knoll rising a few feet above oonieelll 
level of roof. Low plants on knoll are Frankenia fruticulosa, with 
few bushes of Nitraria holding sand on crest. In foreground tus- 
socks of Stipa teretifolia, also Salsola and Lepidiwm foliosum. — 
Several burrows of mutton birds are visible in soft sand below the 
knoll. 

Fig. 2. General view on roof looking west from the knoll 
seen in previous figure. In immediate foreground Frankenia — 
fruticulosa on travertine, beyond the open association on sand, 
Stipa teretifolia, Lepidium foliosum, and Salsola. This association 

alternates with Rhagodia crassifolia shrubland away to horizon. 


. 
(tthe) tas 


PLATE X. 3 


Fig. 1. Cliff vegetation on north coast Eastern Franklin. 
Olearia axillaris on left. Low bushes of Nitraria, Myoporum, 
Threlkeldia, and Frankema pauciflora. 


Fig. 2. “Cove on south coast Western Franklin. Shows in 
foreground Frankema fruticulosa on travertine slope which changes 
abruptly to Rhagodia crassifolia in middle distance. These two 
associations come to the water’s edge on the sheltered side of the 
cove. Facing, in background, is a cliff slope. with Mesembryan- 
themum australe. 


Prats XI. 


Fig. 1. Rhagodia. crassifolia shrubland on roof of Wes- 
tern Franklin. A travertine ridge, with its lighter-coloured flora 
of Frankenia fruticulosa, may be seen running across the field in 
the middle distance. 

Fig. 2. Recent blow-out exposing travertine pavement in 
foreground. The vegetation beyond is of the unstable type on 
sand. Salsola kali very abundant with Stipa teretifolia and 
Lepidium folioswm. Some bushes of Nitraria. The pavement in 
the foreground is said to have been covered with a dense thicket 
of Lavatera plebeja thirteen months before. The dead stems of 
this were very abundant on the loose sand. Photograph taken on 
north coast Western Franklin, looking east. The strait separ ating 
the two islands and the cliffs ‘of Hastera Franklin are seen in the 
distance. 


Fay wl a Ae, 


207 


AN INVESTIGATION OF THE ESSENTIAL OIL FROM 
EUCALYPTUS CNEORIFOLIA, D.C. mag 
(THE “NARROW LEAF MALLEE’’ OF KANGAROO ISLAND.) 


By Puitip A. Berry, B.Sc. 
(Communicated by Professor E. H. Rennie, D.Sc.) © 


[Read August 10, 1922.] 


The principal constituent of this oil is cineol, while it is 
well known that terpenes, aldehydes, and phenols are present. 
The object of this investigation was to determine the average 
cineol content, and the more precise nature and amount of 
these other bodies with which the cineol is associated. 


The crude oil used in this investigation was obtained by 

steam distillation of fresh leaves and twigs, collected at Cygnet 
River, Kangaroo Island, in the beginning of January, 1920, 
from leaf country which had been previously cut about three 
years before. The yield of crude oil was 1 per cent. The 
sample was an orange-brown colour and gave the follow- 
ing constants: —Specific gravity at 12° C.=0°9102; specific 
rotation, (a)p= —10°40; refractive index at 20° C.=1°4707; 
dispersion, 0°01029. The oil was soluble in 1°33 volumes of 
70 per cent. alcohol (by weight) at 20° C. The saponification 
number for the esters and free acids was 7'0. 
Another sample of oil was distilled at Cygnet River, 
_ about the middle of May, 1921, from the same species, in a 
similar stage of growth, and under conditions similar to those 
existing in the above distillation. This oil gave the following 
constants :-—Specific gravity at 13° C.=0°9248; specific rota- 
tion, (a)pD=—4°91°; refractive index at 20° C.=1°4670; 
dispersion, 0°00979. The oil was soluble in 1:05 volumes of 
70 per cent. alcohol at 20° C. 

This second distillation was performed to obtain an idea 
of the difference between the oils distilled in the summer and 
in the winter. The samples cannot be considered strictly 
comparative, however, since, in the first case, the distillation 
' was continued until the leaf was exhausted of oil, while the 
second sample was distilled under ordinary commercial con- 
ditions, that is to say, until the amount of oil distilling was 
very small in comparison to the water, and in consequence 
contained less than the previous sample of the higher boiling 
or less volatile constittients of the leaf. 


208 


Physical Constants.—In order to save repetition and 
avoid ambiguity, the following explanatory note is inserted. 


Specific Gravity.—The specific gravity was taken with as 
large a pyknometer as the amount of liquid permitted. The 
pyknometers were standardized with water at 15° C., and 
when the specific gravity was taken at another temperature, 
the result was calculated to +2°C. by using a coefficient of 
cubical expansion of 0°00075 for each ° C. The specific gravity 
refers to that calculated for {|S° C., except where other tem- 
peratures are given. 

Rotation.—The rotation refers to the actual rotation in a 
100 mm. tube. 

Refractive Index.—The refractive index was taken with 
an Abbé refractometer, and the result calculated for a tem- 
perature of 20° C. by adding or subtracting 0°00047 for each 
°C. by which the temperature exceeds or falls short of 20° C. 
The refractive index scale of the instrument is so arranged 
that it reads directly the refractive index for the mean of 
the D lines of sodium light. 

Dispersion.—The dispersion figures given refer to the dis- 
persion between the C and F lines of hydrogen (656°3 uu. to 
4861 uu.).. The dispersions were taken at the same tempera- 
ture as the refractive index, but were not calculated for 
20° C., as the correction over a small range of temperature is 
negligible. 

Temperature.—All thermometer readings have been cor- 
rected for the unimmersed portion of the stem of the 
thermometer. 

Eaperimental.—The first sample of oil distilled was the 
one used throughout in this investigation. 


A. DISTILLATIONS. 

A 1.—The oil was first subjected to dry distillation. On 
distilling the crude oil, it commenced boiling at 80° C., and 
some acid, water, and volatile aldehydes distilled over first. 
As the oil appears to suffer decomposition by prolonged heat- 
ing at a high temperature, the quantity used for distillation 
was 80 ccs., since this small quantity could be distilled quickly 
and at the same time represented the minimum required for 
tests. The following results are the average of four dis- 
tillations : — 


Temperature. Amount. Rotation. 
Below ‘l79rs?-C2 4 ue 7% ie € 
17h? CIBER 47ers of ot (655 — 7° 
186° C.-207° C. aig bes fos ok tee —10° 
207? et koe re yah Bie — 23° 


ad 
{ 
2 m 


, 


209 


By slow distillation (4 ccs. per minute), the first fraction 
was found to have a dextro-rotation of 1°. This fraction con- 
tained a large quantity of aldehydes, their presence being 
proved by Schiff’s reagent. Aldehydes were also found in 
quantity in the last fraction. Distillations under reduced 
pressure gave very similar results. 


The oil was next subjected to steam distillation. 
A 2.—One litre of the crude oil was steam distilled in 


seven hours. The distillate was collected in the following 


fractions : — 

Amount. Rotation. 
(a)re..-- 00 CGS. Pe TOP 
tbe OU Ges. a Be 
‘(ey *... 100" ces. — 76° 
(d) ... 100 ces. — 638° 
fedin ¢ oJ 00) ces: — 6:0° 
prt... FOO (acs: care ad le 
ko) 235 AO cess: — 70° 
(h)~ ... 100 ees. — 84° 
Gyo) 100"ees: 2a — 89° 
(j) ... 100 ces. ‘A —11°7° 
Gyo hee» DOL CES. aa — 29°8° 


A 3.—Another litre of the crude oil was similarly steam 
distilled in seven hours and the distillate collected in the fol- 


lowing fractions : — 
Specific Refractive Disper- 


Amount. Rotation. Gravity. Index. sion. 
(a) 25 ces. — 5°56° — — — 
(b) 25 ccs. — § 74° 0°9105 14690 0°01003 
(c) 788 ccs. — 69° 0°9047 1°4682 0°01001 
(d) 78 ces. —12°2° 0°9210 A737 0°01043 
(e) 42 ccs. —26°15° ee hin =e 


In the next three distillations the oil. was treated with 
sodium hydroxide either before and/or during distillation, the 
object being to fix, and thus ensure as far as possible the 
removal of the aldehydes. : 

A 4.—One litre of the crude oil was shaken with 300 ccs, 
of a 5 per cent. sodium hydroxide solution. The oil was 
separated three days later and steam distilled. The distillate 
was collected in the following fractions :— 


Specific Refractive Disper- 


Amount. Rotation. Gravity. Index. sion. 
(a) 63 ccs. — 519° 0°9263 1°4702 0°00996 
(b) 786 ccs. 2)-6°72 0°9049 1°4679 0°01012 
(c) 46 ccs. — 9°85° 0°9177 - 1°4712 0°01060 
(d) 68 ces. —26°18° 0°9475 1°4875 0°01204 


A 5.—One litre of crude oil was shaken with 600 ccs. of 
a 20 per cent. sodium hydroxide solution for several days and 


- 


210 


steam distilled without separating the alkali, The distilla- — 


tion. took. about.two and a half hours:— | 
- Specific Refuoctine Disporsl 


Amount. Rotation. Gravity. Index. J? pen: 
(a) 28 ces. ord tek 0°9004 1°4669 0°01009 
(b) 815 ces. —519°° Pr 00aea vee AGT ae 0°01011 
(c) 50 ccs. —5°64° 0°9257 1°4759 * (0°01087 


A 6.—One litre of crude oil, stored in the dark since dis- 
tillation, was shaken with 600 ccs. of a 16 per cent. sodium 
hydroxide solution and steam distilled in the presence of the 
alkali. The rotation of the oil was oy 28°. The distillate 
was collected in two fractions : — 


Amount. Rotation. — Specific gravity. 
(a) 750 ees. —4°84° 0°9065 
(b) 150 ces. — 4°8° iridae 0°9144 


B. EstiMaTIon oF CINEOL IN CRUDE OIL. 


It was found that the resorcinol. method of estimating 
cineol gave very erroneous results with this oil.. The aldehyde 
and other bodies present were, to a large extent, extracted by 
the resorcinol solution, thus giving results which were far too 
high. In one test, the oil which was not absorbed by the 
resorcinol solution had the same rotation as the crude oil 
itself, indicating that the resorcinol absorbed other bodies 
besides cineol. This is confirmed by the appreciable rota- 
tion (often as high as —2) of the cineol separated from the 
cineol resorcinol compound. 

Phosphoric Acid Method.—In connection with the estim- 
ation of cineol by the phosphoric acid method, the writer 
carried out estimations on control samples, containing various 
proportions of pure cineol diluted with ordinary commercial 
turpentine. 

Those samples containing about 70 per cent. cineol (by 
volume) gave excellent results, while 80 pei cent. cineol 
samples were slightly low, and 60 per cent. cineol and less 
gave very erroneous results, a 60 per cent. sample only aver- 
aging 45 per cent. cineol and a 50 per cent: one appeared to 
contain 30 per cent. cineol. It was observed that accurate 
results were only obtained when the cineol phosphate mass 
(before pressing) was of a powdery nature, a pasty mass of 
cineol phosphate invariably giving low results. A probable 
explanation of these low results is that the pasty condition is 
caused by the solvent action of the other constituents on the 
cineol phosphate compound. The addition compound so dis- 
‘solved being removed on pressing the solid cake thus causes 
a low result. A pasty compound of cineol phosphate resulted 


ew tak. 


+> 


2 ‘ 


on mixing the crude oil with phosphoric acid, and this pasty 
nature of the phosphate compound could not be.overcome even 
by the addition of an equal volume of pure -cineol to the crude 
oil. When, however, the crude oil was refined by steam distilla- 
tion by removing from 5 to 10 per cent. of the higher boiling 
fractions, the cineol content estimated by this method,. and 
calculated for the crude oil, was equal to from 60 to 63 per 
cent. cineol. 
Even these results are peel somewhat low, as the 
addition compound with phosphoric acid was still slightly 
_ pasty. From other estimations and comparisons made at the 
time, it was concluded that a more correct figure was 
| from 65 to 68 per cent. cineol. 
Arsenic Acid Method.—The arsenic acid method, as pro- 
_ posed by Turner and Holmes in America in 1914, was also 
tried with the same. control samples as were used above, and 
although in some cases good results were obtained, it does: 
_ not generally appear more accurate than the phosphoric acid 
| method, when working on this oil. It furthermore has the 
drawback that a powdery addition compound is not obtained, 
and thus no indication is given as to the probable accuracy 
of a particular estimation. 


oad 


C. THE SEPARATION OF TERPENES AND OTHER HYDROCARBONS 
UNABSORBED BY RESORCINOL SOLUTION. 


The formation of a water soluble compound of cineol and 
resorcinol formed a ready and convenient method for separ- 
ating cineol from the hydrocarbons in the oil. The large 
middle fractions of the steam distillations, consisting almost 
entirely of cineol and hydrocarbon bodies, were severally used 
for this separation. They were shaken repeatedly with a 
50 per cent. resorcinol (aqueous) solution to remove the cineol. 

_ Other oxygenated bodies, such as aldehydes, etc., were also 
largely removed by this treatment. The oil which was not 
absorbed by the resorcinol solution was steam ‘distilled, and 
from several estimations was — to constitute about 20 pec 
cent. of the crude oil. 

Fifteen ccs. of one of these fractions, which was not 
absorbed by resorcinol and which had a rotation of —13°5°, 
were distilled and the distillate eolteered in the following 


fractions : — 
Return iize, ae EER IAS Amotint: Rotation. 
Pada pr (ae CoL76% Cpe, yt 64 ces. ~ —12°48° . 
+ -(b) 176° G.-1772.C.. 5 Jig Ay esi ¢ie Vp l2-989 
(c) 177° C.-195° C. 34 ccs. —12°98° 


The greater part of the last aie had distilled at 
180° C. 


212 


Oxidation. with. Niirin-Adid Beverel>-ces) aa 


fraction were oxidised with dilute nitric acid (10 per cent.). : 
From the oxidised product two acids were separated; — 


terephthalic acid, which was identified by its insolubility in 
water, alcohol, ether, benzene, etc., and by its subliming on 
heating; and p-toluic acid, which was soluble in alcohol 
and melted constantly at 179°C. The formation of these 
acids indicated the probable presence of cymene. The middle 
fraction of steam distillation A 5 was treated repeatedly with 
a solution of resorcinol (50 per cent.). The oil unabsorbed 
by resorcinol solution had a rotation of —11'2, and was 
shaken with a solution of sodium bisulphite (35 per cent.) to 
remove any aldehydes which had not been polymerised by 
the alkali treatment. The oil was separated and steam dis- 
tilled. The specific gravity was 0°861 and the rotation 
—10°4°, showing that the bisulphite treatment had removed 
some laevo-rotatory aldehyde. 

Schiff’s reagent gave no immediate coloration, but a faint 
violet. developed on standing. 


The sample was distilled and collected in the following © 


fractions : — 
Rota- Specific Refract. 
Temperature. Amount. tion. Gravity. Index. 
(a) 175°-178° C. 21 ccs. — 9°38° 0°8605 1°4820 
(b) 178°-181° C. 16 ccs. —10°14° — 0°8619 1°4827 
(c) 181°-186° C. 13 ces. — 92° — whe 
(d) Residue — — 17° a = 


On shaking the fraction marked (b) with a mixture of 
four volumes of concentrated sulphuric acid and one volume 
of water, allowing to stand for 24 hours separating, and 
steam distilling, and repeating the whole process until no 
further charring occurred, the residue so obtained had the 
following constants:—Rotation, +0°16°; refractive index at 
20. ee 
approximate to those ascribed to cymene. . 

Separation of Pure Cymene.—In order to separate pure 
cymene, the oil unabsorbed by resorcinol solution from the 
middle fraction of steam distillation A5 was treated as 
follows : — { 

It was shaken with a solution of sodium hydroxide (15 
per cent.), the oil separated and steam distilled in the presence 


of caustic soda (digested with solid caustic soda) for several” 


hours, then heated for half a day with sodium metal and 
again digested with solid caustic soda for one day. After 
again steam distilling in the presence of caustic soda and 
digesting with sodium metal and solid caustic soda several 
times it was again distilled. The oil distilled between the 


1°4936; ‘dispersion, 0°01374. These figures closely 


213 


ad 1615.0 but the bulk distilled. from 


175-178° C. The distillate had a rotation of —3°4°; refrac- 
tive index, 14814; and dispersion, 0°01294. No violet colour 
was formed on standing with Schiff’s reagent for 20 minutes. 
The resulting impure cymene was shaken with four successive 
quantities of Beckman’s chromic acid mixture and steam 


distilled. It was then shaken with potassium permanganate 


solution in the cold, steam distilled, and collected in two 
fractions. ‘The first fraction had an odour strongly reminis- 
cent of cineol, while that of the second fraction was more like 
cymene. This latter fraction was digested with sodium for 
a day, distilled and collected in two fractions : — 


Rotation. 
eso 7 8oC. et +1°48° 
(b) 178°-180° C. nt —(0°44° 


The first fraction (a) was shaken with excess of potassium 
permanganate solution in the cold and steam distilled. The 
distillate, which had a rotation of —0°26° and a refractive 
index of 1°4872, was digested over sodium metal and again 
distilled, when it boiled constantly at 177° C.; after drying, 
it gave the following results on combustion :—Carbon, 88°05 
per cent. ; hydrogen, 10°38 per cent. Theoretical for cymene: 
carbon, 89°48 per cent.; hydrogen, 10°52 per cent. 

A portion of the same sample on oxidation with hot 
potassium permanganate solution, as recommended by Wal- 
lach, yielded parahydroxy isopropyl benzoic acid, which on 
repeated crystallization melted at 158° C. Another separa- 
tion of cymene and terpenes from steam distillation A 6 (a), 
by means of resorcinol solution gave a sample with a rotation 
of —12:05° and a specific gravity of 0°8536. After repeated 
digestion with sodium metal, 128 ccs. were distilled and col- 
lected as follows : — 

Rota- Specific Refract. Disper- 


Temperature. Amount. tion. Gravity. Index. _ sion. 
(a) Below 175° C. disces:.°|— 36° (2), . 08567 T4777 \- OF OL255 
(b) 175°-178° C. J0 Ces Oibee O8on2., 4/95. 001288 
fe) 178°-181° C. 26° Secs. 116°. 08509" 14821 = 00138438 
(d) 181°-184° C. AS5ecs. —10°5° — 1°4832 0°01344 


The rotation of the above fractions is almost certainly 
due to laevo-rotatory terpenes. 


Limonene.—Although these fractions contain cymene, 
there is little doubt, judging from the boiling point, specific 
gravity, and other characteristics, that limonene is also pre- 
sent, and that the laevo-rotation of the above samples is due 
to this terpene. 


214 


Pinene.—It is also very probable that dextro-pinene is 


present in small quantities, as 7 per cent. of the oil distils 
below 175°5° C., and by shaking out with resorcinol solution, 
a small amount of dextro-rotatory body was obtained, with — 


an odour characteristic of pinene. Attempts to separate the 
characteristic nitrosochloride were not successful, but this 
does not prove the absence of pinene, as it is very difficult 
to prepare the nitrosochloride from a high-rotation pinene. 


D. ALDEHYDES. 
Estimation.—The total amount of aldehydes occurring 
in the crude oil was estimated by shaking 20 ccs. of the oil 
in a cassia flask with about 150 ccs. of a 35 per cent. sodium 


bisulphite solution ‘and heating on a water bath for three 


hours. The unabsorbed oil was brought into the neck of the 
flask by the addition of more bisulphite solution and the 
volume read off when cold. By this method, a figure of 
75 per cent. of aldehydes was obtained. An identical value 
was also obtained on a sample of oil distilled in May of the 
following year. This would indicate that the aldehyde con- 
tent does not vary appreciably with the season. 


Separation of the Aldehydes from the Orl.—It was found 
that the oil contains more than one aldehyde, and they were 
first separated from the higher boiling fractions of the oil 
by shaking the fraction with twice its volume of sodium 
bisulphite solution (35 per cent.). By this method, a large 
quantity of bisulphite addition. compound was precipitated, 
which was filtered on the Buchner funnel. The filtrate con- 
sisted of oil and aqueous liquor, and these were separated, 
and the aqueous portion used to wash the solid residue in 
the funnel. This washing was continued until no more oil 
appeared in the filtrate. The residue was then washed with 
fresh sodium bisulphite solution, followed by a washing with 
alcohol-ether mixture. The solid cake was dried, decomposed 
with hot sodium carbonate solution, and the aldehyde 
separated. This aldehyde is here referred to as aldehyde A.‘ 


On treating the liquor separated from aldehyde A (containing 


sodium bisulphite, sodium carbonate, etc.) with caustic soda 
solution, a further separation of aldehyde took place. This 
aldehyde, called aldehyde B, was extracted with benzene, 
which was later evaporated off. The aqueous portion of the 
filtrate from the separation of the solid residue was treated 
with hot sodium carbonate solution, but no separation of 
aldehyde occurred. On the addition of caustic soda solution, 


(1) The aldehydes separated are here indicated by the letters 
A, B, and C, as it is later shown that they are mixtures. 


a 
- e 
{ ba 
_ 
Zo t 


oO 


215 


however, a large amount of aldehyde separated, which was 
extracted with benzene, the latter being evaporated off. This 


constitutes aldehyde C. The oil which was not absorbed by 


the sodium bisulphite contains cineol, cymene, terpenes, and 
sesquiterpenes, and was used for ‘the separation of the 
sesquiterpene. This separation of aldehydes was performed 
on the last fractions of steam distillations A 3 and A 4, and 
also on the final fractions of a steam distillation of 2 litres of 
the crude oil. These latter are indicated by the letters 
e and d in the table which follows (Table 1). The results 
tabulated below are calculated for 100 ccs. of oil used. 


TABLE 1. 
. 

Sample Aldehyde A Aldehyde B a Aldehyde C | Unabsorbed Oil 

| Amt. Amt. Be mt. 
No. |Distl.|/Fr’ct.| Rotat.| in | Rotat.| in | Rotat. . “in Rotat. in Rotat. 

| ces. ces. | ees. | ccs. 

; | 
a A3 e —26°15 | 14°2 | —50°0 3°9 | —88°5 66 | —79°3 50°0 | = 

b A‘4 i-d —26°18 | 16°0 | —53°6 2°2 | —86'8 7°8 | —74°3 500 | —83 
¢ |P’n’lt|imate| 24-68 133 | —43-9| 3:9 | —90°0| 4:3 | —741| 586 | —15°1 
a | 164 —54°9 | 4°5 ae 3°6 = 5L5 : —9.5 


final | —25°7 


As the yield of the separate aldehydes was so small when 
working on these fractions, a quantity of the higher boiling 
fractions (called the residual oil) which remain in the still 
after refining the oil on a commercial scale was obtained 
from Kangaroo Island. This residual oil was obtained from 
Eucalyptus cneorifolia only. . 

The oil was steam distilled, and the distillate had a 
rotation of —27°88°. Several separations as described above 
were performed on this oil, and the results tabulated are 
calculated for 100 ces. used. 


Tanne 2. 
Aldehyde A _ _ Aldehyde B | Aldehyde C | Unabsorbed Oil 
No. | Pe ee ee 
Amount Amount ieee Amount | 
| in ees, Rotation Fai Boa | Rotation | Tale [Rotation | | ane | Rotation 
i 1 
e 16°90 | —20°65 2°2 — Hes —60°0 | 38°2 —4°15 
f 117° ~«|«~—15'8 07 = 25°0 —62°7 36°0 —2°31 
£ 11-4 | —12°75 | sy aa 27-3 —63 8 | 324 — 2-55 
} | | 


The semicarbozone prepared _ from aldehyde A (Experi- 
ment g) melted constantly at 208°C. That prepared from 
aldehyde B (Experiment g) melted constantly at 199° C. 


The various aldehyde. bemoans separated were then 
examined individually. 


216 


Aldehyde A. 3 , 


Aldehyde A separated from the crude oil in Experi- | 
ment c (see Table 1) had the following constants:—Refractive | 
index at 20° C.=1°5110; dispersion, 0°01685. 

As the amount of aldehyde separated from the crude 
oil fractions was very small, the aldehyde A from Experiments 
a, b, and d were mixed. These mixed aldehydes gave the 
following constants:—Specific gravity at 17°7° C.=0°9724; 
corrected for +12 ° C.=0°9744; rotation, -—52° refractive 
index at 20° C.=1°5096; dispersion, 0°01673. . 7] 

Aldehyde A separated from the residual oil (Experi- 
ment g, Table 2) gave the following constants :—Specifie 
gravity at 12°C. =0°9720; corrected for 12° C.=0°9742; rota-_ 
tion, —12°75°; refractive index at 20° C.=1°5203; disper-— 
sion, 0°01876. 

Preparation of the Oxime.—5 ccs. of the latter sample 
were dissolved in 10 ccs. absolute alcohol, and to the solu-— 
tion were added 10 ccs. of a saturated aqueous solution of 
hydroxylamine hydrochloride. The mixture was then made 
alkaline with sodium carbonate solution and heated on the ~ 
water bath for five hours, and then poured into cold water. — 
The oxime should crystallize under these conditions. It was 
found that by changing the water into which the heated mix- — 
ture was poured, the crystallization of the oxime was 
facilitated. Although, in this instance, the water was changed 
several times, great difficulty was experienced in obtaining — 
the oxime in a crystallized condition. : 

After crystallizing by this method and recrystallizing from 
alcohol, the oxime melted constantly at 57:2° C. Attempts 
to obtain a crystalline oxime from the aldehyde A samples 
having a high rotation always proved unsuccessful, as a pasty 
mass always resulted on pouring the heated liquor into water. 
A sample of this aldehyde A having a high rotation (—44°) | 
was recombined with sodium bisulphite, filtered, well washed | 
with alcohol-ether mixture, and decomposed with hot sodium 
carbonate solution. By this treatment, one-half of the alde- 
hyde was lost, and the rotation of that separated had 
diminished to —14°5°. The specific gravity was 0°977. As 
the loss of aldehyde on recombining with sodium bisulphite 
seenied inordinately great experiments were performed to 
account for the loss, and it was found that the washing with 
alcoholic-ether mixture dissolved a considerable amount of 
the bisulphite compound. These alcohol-ether washings were 
evaporated to a low bulk, and decomposed with a solution of 
sodium carbonate. The aldehyde thus separated had a high 
rotation (—44°). It therefore appears highly probable that 


ym 


~o a 


‘ 
; 
MB core aor ae rte 


217 


aldehyde A consists of two aldehydes, and that the bisulphite 
addition compound of the one with the high rotation is more 
soluble in alcohol-ether than the other. Another sample of - 
aldehyde A having a rotation of —34°, after recombining - 
with sodium bisulphite, filtering, washing, and decomposing 
with sodium carbonate, had a rotation of —5°9°. The oxime 
was also prepared from this latter sample, and it easily 
crystallized when poured into water. It melted constantly 
at 55°7° C. It was noticed that the less the rotation, the 
easier the preparation of the oxime became. 

Conclusion.—The melting point of the oxime of cumic 
aldehyde is 58° C., and as the constants of the aldehyde with 
the small rotation are very like those of cumic aldehyde, and 
the melting points of the oximes are approximately the same, 
it appears certain that cumic aldehyde is a constituent of this 
oil. The other portion of this aldehyde A is probably the 
aldehyde which H. G. Smith previously named aromadendral. 
(This term has recently been extended to cover the whole 
of the aldehydes occurring in Eucalyptus oils.) 


Aldehydes B and C. 


The additive bisulphite compounds of these aldehydes 
were not decomposed by sodium carbonate, but were by caustic 
soda. Aldehyde C consists of two aldehydes, and it seems 
probable that aldehyde B above is a mixture (probably in 
different proportions) of the same two aldehydes which’ con- 
stitute aldehyde C. It is probable that the addition com- 
pound of these two aldehydes is very soluble, and hence while 
the bulk of it is soluble in the sodium bisulphite solu- 
tion, a small amount is precipitated along with the cumic 
aldehyde and the aromadendral. The aldehyde B separated 
by Experiment g from the residual oil was distilled under a 
pressure of 30 mm. The temperature rose slowly during the 
distillation from 132° to 141° C. The distillate gave the fol- 
lowing constants:—Specific gravity at 72° C.=0°9469; cor- 
rected to 15° C.=0°9502; rotation, —99°7°; refractive index 
at 20° C.=1°4899; dispersion, 0°01341. 

Aldehyde C separated from the crude oil by Experi- 
ment c had a refractive index of 1505. The aldehydes con- 
stituting aldehyde C were separated from each other by 
means of neutral sodium sulphite solution. 

Method.—The aldehyde was shaken with a solution of 
crystallized sodium sulphite (35 per cent., neutralized to 
phenolphthalein), the liberated sodium hydroxide being con- 
tinuously neutralized with normal sulphuric acid. The un- 


combined aldehyde was extracted by benzene. The 


G 


218 


combined aldehyde was liberated by caustic soda solution and — 
extracted with benzene. This separation was performed on — 


aldehyde C obtained from the residual oil by Experiment e. 
The aldehyde which combined with the neutral sodium sul- 
phite, after separation by the above process and evaporation 
of the benzene, was distilled under reduced: pressure. The 
bulk distilled at 121° C. at a pressure of 25 mm. and was 


quite colourless. The following constants were obtained :— 


Specific gravity at 18° C.=0°9442; corrected for 15° C.= 
0°9462; rotation, = —65°61°; refractive index at 20° C.= 
14834; dispersion, 0°01247. 


The aldehyde which did not combine with neutral sodium 


sulphite, and which was extracted with benzene, was 


separated from the latter and distilled under reduced pressure. 


It did not distil at so constant a temperature as the sample 


above. The temperature rose from 125 to 135° C. during the 
distillation, while the pressure decreased from 28°5 to 
20°55 mm. This aldehyde had a rotation of —70°64°, and 
was not oxidised by exposure in the atmosphere, the rotation 
remaining unaltered after exposure in an open beaker for 
fourteen days. This point is here stressed, as H. G. Smith,@ 
in his separation of these aldehydes from Eucalyptus oils, 
states that this fourth aldehyde is oxidised by the air and 
closely resembles the phellandral separated by Schimmel and 
_Co. from water fennel oil. (The volatile oils, by Gildemeister 
and Hoffman, page 432.) The aldehyde here separated can- 
not be phellandral, as the latter is oxidised on exposure to 
the air. This separation was also performed on aldehyde C 
of Experiment g. The aldehyde which combined with the 
sodium sulphite after separation was distilled under reduced 
pressure; about two-thirds of it distilled at 126°C., the 
pressure being constant at 32 mm.; the temperature then 
gradually rose to 130°C. The distillate gave constants prac- 
tically identical with those from Experiment e (see above). 


These constants are practically the same as those obtained 
in the first separation and agree with those given by H. G. 
Smith for Cryptal. No crystalline oxime could be separated 
from this aldehyde. 

The aldehyde which did not combine with sodium sul- 
phite was distilled under reduced pressure and collected in 
two fractions : — 


(a) 1219-126° C, 21 mm. 33 ccs. 
(b) 186°-140° C. 32 mm. 26 ccs. 


(2) ‘A Research on the Eucalypts and their Essential Oils,’’ by 
R. T. Baker and H. G. Smith, p. 286. 


(3) Ibid, p. 387. 


219 


About 20 ccs. of (b) distilled between 136 and 138°C., 
and then the temperature rose slowly to 140°C. The follow- 
ing constants were obtained : — . 


Sample a. Sample s. 
Ep ciic gravity oes ia) JOreovo 0°9553 
Rotation . —67°92° — 72°99° 
Refractive index at 90° C. ... 1°4888 1°4921 
Dispersion any : . 001825 0°01363 


The preparation of the oxime was tried on these samples, 
but the product was only crystallized with great difficulty. 
By carefully crystallizing from chloroform, however, minute 
crystals were obtained ; one crystal grew sufficiently to permit 
of its removal by mechanical means. When washed and dried 
it melted at 84-85°5° C. As this aldehyde is not identical 
with any yet separated from Eucalyptus oil, it is proposed 
to call it cneoral. The amount available did not permit of 
any extensive work being performed on it, and it is possible 
that it may later be proved to be identical with some aldehyde 
already separated from some other Eucalyptus oil. 


Conclusion.—This oil appears to contain four aldehydes— 
cuminal, aromadendral, cryptal, and a fourth one, which is 
here called cneoral. The first three of these agree with the 
aldehydes separated by H. G. Smith (loc. cit.) from 
Eucalyptus oils. | 

EK. SESQUITERPENE. 


The last fraction in the distillation of the oil contained 
a sesquiterpene, which answered to H. G. Smith’s tests for 
aromadendrene. The oil which was left unabsorbed after 
shaking the higher boiling fractions of the crude oil with a 
35 per cent. sodium bisulphite solution (to remove aldehydes) 
was equivalent to 4 per cent. of the crude oil and had a 
rotation of —15°1°. As the quantity of sesquiterpene pre- 
sent was small, the residual oil left after refining the crude 
oil on a commercial scale was shaken with a solution of sodium 
bisulphite (35 per cent.). About one-third of the oil was not 
absorbed by this treatment, and it gave the following con- 
stants:—Specific gravity =0°9421; rotation = — 2°55°; refrac- 
tive index=1'4800; dispersion, 0°01086. This sample was 
shaken with a solution of resorcinol (50 per cent.) to remove 
_ the cineol, and then steam distilled. The resulting oil, which 
constituted 69 per cent. of the previous fraction, was prac- 
tically colourless and gave the following constants :—Specific 
gravity =0°9356; rotation = —2°7°; refractive index at 20° C. 
=1°4788; dispersion, 0°01055. 

Conelusion.—This oil contains a small percentage of a 
sesquiterpene (not more than about 2 per cent.), which appears 
to be aromadendrene. 

G2 


220 


F. PHENOLS. 


The crude oil contains a small amount of phenols. One ~ 
litre of the crude oil was shaken with 300 ccs. of a 5 per cent. © 
sodium hydroxide solution, and the aqueous liquor separated 
and shaken with ether to remove adhering oil. It was next 
acidified, extracted with ether, and the latter evaporated 
off. The residue was washed with sodium bicarbonate solu- 
tion, extracted with ether and evaporated. The final yield 
was 2 ccs., which is equivalent to 0°2 per cent. of the crude 
oil. Beyond applying qualitative tests, no further work was 
done, owing to the small amount of material available. 


G. ALTERATION OF THE SPECIFIC Gravity, RoTaTion, 
REFRACTIVE INDEX OF THE CRUDE OIL ON KEEPING. 


The following table gives a summary of the results 
obtained : — 
Rota- Specific Refract. Disper- 


Sample. tion. Gravity. Index. sion. 
1. Crude ae Eee dis- 
tilled (Jan., 1920) —9°5° 0°9102 — — 


2. Crude oil stored in a 

tightly-corked bottle 

in the dark for 13 

years —90° 0°9105 = — 
3. Crude oil stored in a 

loosely-corked iron 

drum for 14 years  —7°38° 0°9145 1°4707 0°01029 
4. Crude oil exposed to 

light in white glass . 

for 14 years a. 4°79 0°9311 1°4728 0°01033 


Conclusion.—The specific gravity increases on keeping, 
while the rotation diminishes. The refractive index and the 
dispersion also increase. These changes are accelerated by 
exposure to light, and are probably caused by the polymeri- 
sation of the terpenes and/or aldehydes. 


SUMMARY. 
The results of this investigation have shown that the oil 


from the leaves and twigs of Hucalyptus cneorifolia distilled 
in January has the following approximate composition : — 


Cineol ous ne ee son 3 se GTi 
Cymene iy id Re ae wt 
Limonene vag iy. me, a's id 5% 
Pinene, ) 4%; A ine 3% 
Aldehydes—Cumic Aldehyde — ne ae 
Aromadendral 7% 
Cryptal 2 
Cneoral any fi eS 
Sesquiterpene ... iat rs at 1% 


Phenols, esters, and acids ae eA. si. Oaews 


is 
ig 
ve 
at 


221 


This work has been carried out as a research subject 
under the David Murray Scholarship for Science, and was 
performed under the direction of Professor Rennie, M.A., 
D.Sc., to whom I wish to acknowledge my indebtedness for 
‘advice and help given. I also desire to thank Professor 
Osborn, D.Sc., for the botanical identification of the species. 
Owing to the prolonged nature of the work and the short 
time available to the investigator for work at the University, 
the main part of the research was performed in the laboratory 
of Messrs. A. M. Bickford & Sons, Ltd., and I desire to thank 
them for the facilities and material placed at my disposal. I 
' further wish to thank Mr. J. Hendry, Ph.C., A.I.C., for the 
'many and helpful suggestions offered throughout the work, 
and also Mr. E. Burgess, of Kangaroo Island, who collected 
. the leaves, distilled and donated the oil used in this research. 


222 


ON THE ARRANGEMENT OF THE STRIATIONS OF . 2 
; 
VOLUNTARY MUSCLE FIBRES IN DOUBLE SPIRALS. ~~ 1 


By O. W. Tizes, M.Sc., 


Department of Zoology, University of Adelaide. 
[Read September 14, 1922.] 


Puate XI. 


While examining the muscles of the larvae and adults of | 
a small parasitic wasp, Vasonia, I noticed, recently, that the 
striations were not disposed transversely, as is supposed to 
occur universally in voluntary muscle, but that they were 
arranged in the form of a double spiral. The structure of 
these muscles will be referred to more fully in a later paper. 

The purpose of this note is to draw attention to the fact 
that the striations of voluntary muscle fibres, which histol- 
ogists are unanimous in regarding as truly transverse, are in 
reality likewise disposed in the form of double spirals. I have 
observed this in the muscle fibres of the crayfish Astacopsis, 
in the leg muscles of a South Australian grass-hopper, and 
in the voluntary muscles of the much studied water-beetle 
Dytiscus. 

Amongst mammals I have observed it in the muscles of 
the rat, the pig, the dog, the rabbit, the mouse, and, finally, 
in human muscle fibres, and its presence in such widely 
separated groups suggests its universal occurrence. 

In a well-stretched fibre this double spiral arrangement 
of the striations is relatively easy to detect, and is shown in 
fig. 3, taken from the muscles of man. In such muscles it is~ 
possible to begin at one end of a fibre, and focussing up 
and down to travel along the spiral. Muscle fibres, however, 
are usually examined in the contracted condition ; under these 
circumstances it is often extraordinarily difficult to detect 
the spiral nature of the striations. Two methods may, how- 
ever, be adopted : — 

(1) If a muscle fibre has been well flattened, the stria~ | 
tions at the sides of the fibre may actually be observed to | 
bend downwards and out of the plane in which the striations 
on the upper portion of the fibre have approximately lain. | 


| 


This condition, taken from a muscle in the dog’s tongue, is 
shown in fig 4. Generally, however, the striations are so 
very close together that it is possible only with the greatest 
difficulty to notice the smal] change in direction taken by — 
the turn of the spiral. 


223 


(2) A more successful way to exhibit the presence of the 
double spiral is to make a camera lucida drawing of the upper 
striations of a muscle fibre; then focussing through to the 

| lower side make a similar drawing on a separate piece of fairly 
transparent paper. Accurate superposition of the two draw- 
ings will reveal the presence of a single spiral. Care is 
required in the interpretation of the result. Only one-quarter 
of a turn of a spiral is visible in one focus of the micro- 
scope; by focussing successively, therefore, upon the upper 
turns and then upon the lower turns, the additional rise or 
drop in the spiral which would occur if the path of the spiral 
in the thickness of the fibre could be observed, is eliminated. 
The single spiral obtained by the superposition of the two 
drawings will therefore indicate the actual presence of a 
double spiral. 

. The objection which may be taken against this inter- 
pretation is that the change in direction of the striations in 
the upper and lower side of the muscle under examination is 
due to a shearing stress, perhaps due to the pressure of the 
cover-glass on the preparation. By no conceivable method of 
distortion, however, can transverse striations be converted 
into real spirals, and the fact that it is possible to travel 
along the spiral in stretched fibres by successively focussing 
up and down along it, eliminates this difficulty. 

Moreover, it is possible, in focussing through a fibre 
suddenly to come to a focus where there is a discontinuity— 
very faint, but still perceptible—between the upper and lower 
portions of the “‘transverse’’ striation, and it is often possible 
(see figs. 1 and 2) to observe at one focus the crossing of 
striations at this point, clearly indicating their spiral nature. 
Neither is definitely in focus, and while it is possible to see 
both at once, neither can be observed sharply. This is to be 
noticed especially clearly at the terminations of fibres, or in 
those places where they are not quite flat, but where the fibre, 
first in focus, bends slightly out. Under these circumstances 
one obtains a partial view along the longitudinal axis of the 
fibre, and can at one focus obtain a view of a partial turn of 
the spiral. 

It is, perhaps, necessary to add that care must be taken 
not to focus along the plane of contact of two superimposed 
fibres; without this precaution a false crossing effect might 


| readily be obtained by the simultaneous indistinct focussing 


of the striations of two fibres. 


For a clear demonstration of the double spiral the longi- 
tudinal body muscles of chalcid wasp larvae may be 
recommended. 


224 


Since writing the above note, I have observed a do ible 
spiral arrangement of the striation in cardiac muscle fibres. 


DESCRIPTION OF PLATE XII. 


Fig. 1. Voluntary Muscle Fibre from leg of Mouse, x about 
500. The middle of the fibre is bent slightly dodnwande and is 
therefore at a different focus from adjacent parts, which are 
approximately surface views. Unstriped areas inca blurred 
focus. The gradual transition from ‘‘transverse’’ to “crossed”? 
striation is clearly shown as the middle of the fibre (x) comes into 
focus. } 

Fig. 2. Muscle Fibre from leg of Mouse, x1000. The focus 
is along the middle of the fibre, and clearly shows crossing of 
striations, being the optical Stoke of focussing the spirals in one 
plane. 


Fig. 3. Human Muscle Fibre, well stretched, x920. The 
fibre is observed at one single focus. The fibre has become 
stretched to such an extent, that it is at times possible to obtain 
a slightly blurred image of top and bottom of fibre simultaneously 
under these circumstances the complete double spiral may be seen. 


Fig. 4. Portion of Muscle Fibre from Tongue of Dog, x 1400. 
The fibre has been considerably flattened, and shows bending down 
of the striations at the sides of the fibre. Note especially, that 
the blendings are in opposite directions, 


ta ~ 
+ ~ 
f 
“aa ¥ 
E; 4 


225 


AUSTRALIAN LEPIDOPTERA OF THE GROUP GEOMETRITES. 


By A. JEFFERIS Turner, M.D., F.E-S. 
[Read September 14, 1922. ] 


Hitherto I have regarded the moths here dealt with as 


_ forming a single family, the Geometridae. Recent study of 


| 
Uv 
ce 
4 
a 


the families belonging to the Noctuoidea (Caradrinina of 
Meyrick) has caused me to revise my opinions. The families 
Syntomidae, Arctiadae, Hypsidae, Nolidae, and Noctuidae, 
though natural and necessary, yet in the structure of their 
more typical and primitive genera are so closely allied, that 
we must reconsider the value of our family groups of other 
sections of the Lepidoptera. There should be a general 
correspondence in the structural value of family characters, 
though a precise equivalence is, of course, impossible. I 
propose, therefore, to regard the Larentiadae, etc., no longer 
as merely subfamilies, but as groups of family rank. This 
was indeed done long since by Mr. Meyrick in his British 
Lepidoptera where he includes them with the Notodontoidae 
and other families in the larger group Notodontina. The 
weak point in this classification, it has seemed to me, is that 
the relationship, that binds together the geometrid families 
into one group, is not expressed, but is lost in the larger 
and looser complex. This difficulty may be avoided, and I 
think its avoidance is necessary for any satisfactory classifica- 


tion, by placing them as a distinct division, the Geometrites, 


in a larger group the Notodontoidea, which I conceive as 
corresponding generally, but not exactly, with Meyrick’s 
Notodontina. 

The first three families I have already revised in former 
publications, but much remains to be added to bring them to 
completeness at the present date. The Oenochromidae I 
have not yet studied in detail; and of the Boarmiadae I have 
published only a partial and incomplete revision. In these 
two families I shall merely describe a small number of new 
forms. 

Fam. LARENTIADAE. 


I give a new key to the Australian genera, in which 
“many of the names differ from those formerly adopted. Mr. 
L. B. Prout informs me that it has been ascertained that the 
names Cidaria, Larentia, etc., of Treitschke were published 
earlier than HYydriomena, Xanthorhoé, etc., of Hubner. He 
thas also helped me much by indicating the European types 


226 


of some of our genera. The following list indicates the 
changes in name now introduced :—Huchoeca, Hb., becomes: 
(1) Crethets, Meyr.; (2) Euchoeca, Hb. Asthena, Hb., 
becomes (1) VPoecilasthena, Warr.; (2) Minoa, Treit. 
Scordylia, Gn., becomes Chaetolopha, Warr. Hucymatoge, 
Hb., Sect. 1, 2, and 3, become Horisme, Hb.; Hucymatoge, 
Hb.; Heeymatoge, Prout. Hydriomena, Hb., Sect. 1, and 
Sect. 2 and 3 together become Huphyia, Hb., and Cidaria, 
Treit. Xanthorhoé, Sect. 1 and 2, become Xanthorhoé, Hb., 
and Larentia, Treit. a 

The family is a large one; the numerous genera are closely 
allied ; and their classification is difficult. It is a group which 
permits of no primary division; all the characters employed 
for generic distinction are of secondary value. For instance, 
the smooth face characteristic of the Asthena group is found 
also in Saurts, which resembles that group in no other char- 
acter, and had, I believe, a quite different origin. Again 
the possession of a single or double areole, though valuable, 
is a secondary character, which has been independently 
developed in many instances. By its use we may separate 
many pairs of genera, which are as closely or more closely 
allied to each other than to anything else. Such pairs are :— 
Euchoeca—Minoa, Tephroclystis—Mnesiloba, Chaetolopha— 
Cidaria, Epirrhoé (Europe)—Huphyia, Asaphodes (New Zea- 
land)—Xanthorhoé, Dasysternea—Dasyuris. Although the 
character is a valuable one, and indeed indispensable, it is 
not certain that the generic distinctions thereby made will 
always be natural; for no reason can be given why this 
modification, unaccompanied by any other, may not have 
arisen independently in different unrelated species of the 
same genus. In two other generic characters, which I con- 
sider valid, even more difficulty presents itself. Of these the 
first is the pectination of the male antennae. ‘This also is a 
secondary character, and separates groups otherwise similar 
or identical in structure, Xanthorhoé from Euphyia, Larentia 
from Cidaria, Asaphodes from Lmrrhoé, Notoreas from 
Dasyuris, Venusia from Huchoeca. In addition to this weak- 
ness there are also intermediate conditions difficult to classify. 
For instance, Meyrick places the European wittata in 
Xanthorhoé, and this may be its natural position, but the male 
antennae cannot be termed pectinate. This difficulty might be 
got over by broadening the definition of the genus, but the 
Australian percrassata and vacuaria (the latter also placed: 
by Meyrick in Xanthorhoé) have the same antennal struc- 
ture, and closely similar is that of strwmosata, while all three 
species appear to fall more naturally under Huphyia. These 
difficulties occur, however, seldom, and greater difficulties in 


22% 


classification would, I believe, arise if we reject antennal 
characters altogether. 
It will be seen that some of the objections so forcibly 


urged by Mr. Meyrick (Trans. N. Z’d. Inst., 1916, p. 248) 


against the generic value of modifications of the discocellulars 
and origin of vein 5 of the hindwings apply also to characters 
which he recognizes as valid. If applied with impartial 
logic, they would destroy his own, and, I believe, any other 
possible classification of the family. It must be admitted 
that here also intermediate forms occur, though rarely, 
but they are not such as should create any real difficulty. 
Vein 5, which is the second median vein, arises normally 
opposite the termination of the upper primary branch of the 
median trachea, which becomes obsolete in the adult wing, 
but its point of termination is often traceable, often situated 
centrally, but often considerably nearer the radius than the 
cubitus. This is the structure in Huphyia, Xanthorhoé, and 
most of the genera of the family. The approximation of 
5 to 6 is often conspicuous, but I do not attach generic 
importance to it, for 5 appears never to rise from above the 
termination of the upper primary branch of the media as it 
does in the Geometridae (sensu stricto). Usually with this 
origin of 5 the discocellulars are straight or nearly so, but 
not always (see for instance HL pirrhoé sociata, Bkh.). In 
many genera such as Cidaria and Larentico a striking modifi- 
cation occurs. In them 5 arises from well below the termina- | 
tion of the upper primary branch of the media, and there is 
a strong bend approximating to a right angle at its point 
of origin. Usually 5 is also strongly approximated to 4 at 
origin, but not always. In muzcrocyma, for instance, it is 
from not much below the middle, but the discocellular is 
strongly bent at the usual point (not straight, as erroneously 


‘stated in my former revision). This structural division as 


thus understood appears clear-cut, and I have not so far met 
with a really doubtful case. Nor do I find that the genera 
defined by it are less natural than those defined by the areole 
or antennal pectination, when considered as a whole. It 
must, however, be admitted that, as Mr. Meyrick points 
out, difficulties occur in the New Zealand fauna. Larentia 
cimeraria is extremely similar to Xanthorhoé plumbea, but 
here the similarity of grey coloration (doubtless protective) 
and very simple pattern is one that might well have been 
independently acquired, and JI think we can here trust 
structure before appearance. The case of A. adons, L. 
beata, and L. benedicta is more difficult. These certainly at 
first sight appear nearly allied, the last two, however, rather 


_ more closely than the first, which, except in colour, is very 


L 


228 


like X. chorica. Here also I am inclined to trust structure 
rather than appearance. The beautiful green coloration, rare 
elsewhere, is not infrequently developed in this family in 
New Zealand, and the pattern, although striking, is a very 
simple modification of that usual in this family. I admit 
that doubt is possible, and this doubt may be strengthened 
by the resemblance between X. nephelias and L. sericodes, 
which I have not seen. It may be that our structural char- 
acter here breaks down, and that we may have to admit that 
our classification is so far imperfect. This I am easily pre- 
pared to do. The question to me appears to be, not whether 


our classification is perfect, but whether, taken as a whole, 


it is better (more natural), if we reject, or if we admit the 
generic value of the character in dispute. 

Although this question cannot be decided by geographical 
distribution, yet that may throw some light on it. As I 
have been able to examine but few of the European species, 
I have asked Mr. L. B. Prout to give me the results of his 
examination of those included under Hydriomena and 
Xanthorhoé by Meyrick in his study of the European fauna 
(Trans. Ent. Soc., 1892, p. 53). Two species with the areole 
simple, species which Meyrick had not been able to examine, 
are omitted, and vitatta has been transferred to Huphya. 
For the New Zealand fauna my material has been less com- 
plete, but through the kindness of Mr. A. Philpott I have 
_ been able to examine 43 species, and have included 10 more 
on the authority of Meyrick or Prout. I have omitted swb- 
ochraria and subrectaria as Australian species, which may be 
natural immigrants into New Zealand, but were probably 
accidentally introduced, and praefectata, which is allied to 
Venusia. I have expressed the result in numbers and per- 


centages : — x 
European Fauna. Australia. New Zealand. 
Cidaria eee 425% 6 65% 0 00% 
Larentia ey) tag 10°6% 9 98% 17, Sa 
Euphyia ei eo 21:9% 64 696% 9 Toi 
Xanthorhoé ... 40 25'°0% 13 «141% 27 = 50°9% 


Very striking are the great development of Cidaria in the 
European fauna, its slight representation in Australia, and its 
absence from New Zealand; almost equally so the great 
development of Huphyia in Australia; while Larentia and 
Xanthorhoé are most developed in New Zealand. 


Kery To GENERA. 
1. Face smooth 
Face more or less rough-scaled, “usually with 
anterior tuft of scales : cca aton 
2. Posterior tibiae with terminal spurs only ... Sauris 
Posterior tibiae with two pairs of spurs ; 


18. 


URE 
20. 


21. 
22. 
23. 
24. 
25. 
26. 


. Areole simple 


Areole double | 


. Areole small, 


on 9, 10, 11 stalked 


229 


Areole large, ti arising from it separately 


Areole well developed 


Areole double 


. Abdomen crested 

Abdomen without crests .. 

. Posterior tibiae with terminal spurs only . 

Posterior tibiae with two pairs of spurs 

. Forewings with 11 running into 12 

Forewings with 11 free ... . 

. Forewings with 11 running into 12 or absent 

Forewings with 11 free 

. Posterior tibiae with terminal spurs only . 
Posterior tibiae with two pairs of ane 

. Thorax smooth beneath Lea) Bh TNE 


Thorax hairy beneath 


. Forewings with 4 and 5 stalked =f 
Forewings with 4 and 5 mtohs separate 
. Areole simple Fe Ay ne en 


. Areole small, 11 stalked with 10 


=, 


. Hindwings with discocellulars bent, 5 from 
below middle ... 

Hindwings with 5 from above middle of cell 
. Areole absent, 7, 8, 9, 10, 11 stalked . , 


Areole large, 11 arising from it it separately 


. Abdomen ‘crested 
Abdomen without crests ... : 

. Hindwings with 5 from middle of cell, 
male with small tornal lobe ... 


in 


Hindwings with 5 approximated to 4 or 6, 
male without tornal lobe .. 
Hindwings with discocellulars angled, 5 from 


below middle 


Hindwings with discocellulars nearly ’ straight, 
5 from above middle EL Ate 

Thorax with a posterior crest . 

Thorax not crested 


Hindwings with eae angled, 5 from 
below middle 


Hindwings with discocellulars n nearly ‘straight, 
5 from above middle, or rarely from middle 


of cell 


Hindwings of male with 4 absent. 
Hindwings of male with 4 pr esent 
Hindwings of male with 6 absent . 


Hindwings of male with 6 present 


Antennae in male ciliated 


Antennae in male pectinate . 


Thorax smooth beneath . 


Thorax hairy beneath 
Antennae in male ciliated 
Antennae in male pectinate 


Hindwings of male with a well- defined spot 


or patch of androconial scales on Adis 


side 


Hindwings of ‘male without androt oconia 


A. 


59 
Cretheis 


Euchoeca 


Poecilasthena 
Minoa 

8. 
Antimmistis 
Symmimetis 


Gym noscelis 
1. 


Chloroclystis 
Tephroclystia 
Microdes 

Weg 
Anomocentris | 
14 


15. 
Dasysternica 
Scotocyma 
Chaetolopha 
iy 

20. 
Mnesiloba 
18. 
Eccymatoge 
19. 

Horisme 
Eucymatoge 
AE 

24. 
Heterochasta 


Polyclysta 
93 


Cidaria 
Larentia 


Melitulias 
Euphyia 


230 


27. Antennae in male with two pas of pectina- 


tions from each joint ... Diploctena 
Antennae of male with one pair of f pectina- 
tions from each joint ... .. ... Xanthorhoé 
28. Antennae in male ciliated .......... ... 'Dasywris 
Antennae in male pectinate ......... .... Notoreas 


SAURIS PEROPHORA, 0. sp. 

mynpopopos, bearing a pouch. 

3, 30 mm. Head olive-green. Palpi 3, second joint 
rough-scaled above and beneath, terminal joint moderately 
long; olive-green, towards base whitish; terminal joint grey, 
extreme apex whitish. Antennae ochreous-grey. Thorax 
olive-green. Abdomen smooth, without tufts; grey, on 
dorsum greenish tinged. Legs greenish-grey; posterior tibiae 
in male normally developed but without spurs, tarsi elongate, 
first tarsal joint as long as tibiae. Forewings elongate- 
triangular, costa moderately arched, apex pointed, termen 
long, bowed, oblique, in male not incised; whitish largely 
suffused with green and dark fuscous, which form markings; 
five narrow transverse fasciae, dark fuscous in middle, green 
towards costa and dorsum, rather ill-defined; first subbasal, 
second at 4, third at 4; fourth from 2 costa, somewhat 
dentate, consisting of several fine parallel lines, at first curved 
outwards, then inwards, and bent outwards to just before 
tornus; fifth similar from 2 costa to tornus, containing a 
squarish fuscous spot above middle; a whitish dentate sub- 
terminal line following fifth fascia; a terminal series of dark- 
fuscous dots on veins; cilia whitish, apices partly fuscous. 
Hindwings and cilia grey; in male with a large basal dorsal 
pouch extending half-way to costa and to tornus, the dorsal 
edge of this pouch forming an erect concave lobe. 

North Queensland: National Park (3,000 ft.), in March; 
one specimen at light. I might have taken more if I had 
not mistaken it for S. hirudinata, which it closely resembles 
in colour, size, and form: In structural characters it is 
altogether different and resembles S. lichenias rather closely, 
but the pouch of the hindwings is much larger, the first 
posterior tarsal joint proportionately longer, and the palpi 
more roughly scaled, with longer terminal joint. 


Gen. CRETHEIS, Meyr. 


Face smooth. Tongue present. Palpi short, slender, 
porrect. Antennae in male simple, shortly ciliated. Thorax 
without crests, not hairy beneath. Forewings with areole 
small, simple; 7, 8, 9, 10, 11 stalked from areole. Hindwings 
with 3 and 4 stalked or separate, 6 and 7 stalked, 12 anas- 
tomosing with cell to # or beyond. Type, C. cymatodes, Meyr. 


ae 


231 


CRETHEIS CYMATODES, Meyr. 
Euchoeca iophrica, Turn. 


I am indebted to Mr. L. B. Prout for pointing out this 
synonymy. Hindwings with 3 and 4 stalked. 
North Queensland: Cairns, Herberton. Also from New 
Hebrides. 
CRETHEIS ATROSTRIGATA, Warr. 


3, 9, 20-25 mm. Head pale ochreous; face ochreous- 
brown. Palpi whitish-ochreous. Antennae pale ochreous; 
ciliations in male $+. Thorax pale ochreous. Abdomen pale 
ochreous with a few fuscous scales on dorsum. Legs whitish- 
ochreous; anterior and middle pairs pale fuscous on dorsal 
surface. Forewings triangular, costa straight, slightly arched 
towards base and apex, apex pointed, termen bowed, oblique; 
pale ochreous, with more or less pale-fuscous suffusion forming 
slender, indistinct, undulating, transverse lines; several of 
these lines form an obscure basal patch; a blackish discal dot 
beneath 2 costa; a slender, undulating, fuscous line from 
mid-costa, at first outwardly curved, then oblique to dorsum 
before middle; this is followed by several less distinct lines, 
which sometimes combine to form a median fascia; subterminal 
and submarginal lines sometimes containing each several 
fuscous dots; sometimes a terminal series of fuscous dots on 
veins extending into dilia, but these are not always developed ; 
cilia pale ochreous. Hindwings with 3 and 4 separate ; termen 
strongly rounded; as forewings. Underside similar but paler 
and more suffused. Variable; southern examples are slightly 
larger than those from Herberton and lack the subterminal 
fuscous dots, but sometimes have a dark-fuscous tornal spot. 

North Queensland: Kuranda, near Cairns, in May; 
Herberton in October, November, December, and January. 
Queensland: Rockhampton, Bundaberg in July, Brisbane in 


_ December, Rosewood in April. 


Gen. PoECILASTHENA, Warr. 


Type P. pulchraria, Dbld. In most of its characters 
this approaches Oporima, Hb., type O. dilutata, Bkh., but I 
do not think there is any really close relationship. O. dilutata 
differs in the peculiar structure of the areole, of which the 
dividing bar (vein 10) arises from the end of the cell, and 
the posterior extremity of the areole is prolonged to reach 
half-way, or nearly half-way, from cell to apex. In the latter 
respect it agrees with the allied genus Operophtera, Hb., 
which, however, has the areole simple. To Poecilasthena I 
refer, with one exception, all the Australian species formerly 
referred to Asthena, Hb. 


232 


POECILASTHENA THALASSIAS, Meyr. 


The male of this species has a very large extrusible tuft 
of fuscous hairs on the underside of the apex of the abdomen. 
This will serve to distinguish it from A. pulchraria; A. 
balioloma, Turn., has also a smaller, stiffer, less woolly tuft 
in the same situation. 


POECILASTHENA STHENOMMATA, Nl. sp. 
oevopparos, strong-eyed. 


3,9,30-32 mm. Head grey, between antennae whihigee 
face fuscous- brown, lower edge whitish. Eyes rounded, in 
female rather large; in male much enlarged, so that a line 
drawn from one outer edge to the other is longer than the 
breadth of the thorax. Palpi in female small, in male 
minute; grey-whitish. Thorax grey mixed with whitish. 
Abdomen whitish with grey irroration. Legs ochreous-whitish. 
Forewings triangular, costa slightly arched, middle portion 
nearly straight, apex acute, termen bowed, oblique, sub- 
dentate; whitish with dull- -creenish markings, thinly scaled ; 
costa with numerous grey spots, which form the commence- 
ment of greenish transverse lines, more or less undulating; a 
basal patch of three or four close-set lines; a median white 
band containing two fine interrupted lines, succeeded by a 
dark-fuscous discal dot beneath mid-costa; beyond this is an 
undulating greenish fascia containing white dots on veins; 
terminal area whitish with two or three undulating, greenish, 
transverse lines; a fine fuscous terminal line interrupted on 
veins; cilia grey-whitish. Hindwings with termen rounded, 
dentate, a stronger acute tooth on vein 4; as forewings, but 
. base whitish. 

The enlarged eyes of the male is a very exceptional 
character. 

North Queensland: Evelyn Scrub, near Herberton, in 
January; three specimens received from Mr. F. P. Dodd. 
New South Wales: Mount Gregson, Liverpool Range, in 
March; one female, in Coll. Lyell. 


POECILASTHENA XYLOCYMA, Meyr. 


New South Wales: Moruya, in October; one female 
specimen corresponding well with a female from Western 
Australia (Waroona) in May, in Coll. Lyell. Also from 
Victoria: Melbourne, Beaconsfield. 


POECILASTHENA PANAPALA, Nn. Sp. 
mavatraAos, all-tender. 


3,24mm.; 9,28mm. Head brownish-grey, anteriorly 
broadly white; face dark fuscous. Palpi whitish; terminal 


—— 


od 


233 


joint dark fuscous. Antennae dark grey, towards base whitish ; 
ciliations in male minute. Thorax brownish-grey. Abdomen 
grey, mixed with whitish; paired fuscous dots on dorsum of 
each segment. Liegs fuscous; posterior pair except tarsi 
whitish on dorsum. Forewings triangular, costa slightly 
arched, apex round-pointed, termen bowed, moderately 
oblique; grey-whitish with numerous, fine, curved, brownish- 
grey, transverse lines and suffusion; a dark-fuscous discal dot 
beneath 2 costa; a slightly darker slender fascia from 2 costa 
to mid-dorsum, edged with wavy darker lines; an interrupted 
fuscous terminal line; cilia brownish-grey, apices paler. Hind- 
wings with termen rounded, slightly wavy, and slightly angled 
on vein 4; as forewings but without discal dot. Underside 
grey, with obscurely darker discal dots on both wings, two 
obscure lines on forewing and three on hindwing towards 


_termen. 


Very near P. xylocyma. The best point of distinction 
in the female appears to be in the terminal line, which does 
not consist of paired dark-fuscous dots. The male has no 
recurved hairs on tornus of hindwings. 

New South Wales: Mount Kosciusko (5,500-6,000 ft.) 
in January, two male specimens; Wentworth Falls, near 
Katoomba, in April, one female in Coll. Lyell. 


Gen. Minoa, Treit. 

Type iM. murmata, Scop., from Europe. This genus 
comes very close to Asthena, Hb., type A. candidata, Schif., 
which differs in having 7, 8, 9, 10, and 11 stalked from areole. 
The stalking of 11 is unusual in the family and appears to be 
a good generic character. Only one Australian species, J/. 
euthecta, Turn., has been recognized. 


Gen. ANTIMIMISTIS, nov. 

dvTitpustis, imitating, modelled after. 

Frons with strong anterior tuft of scales. Tongue pre- 
sent. Palpi rather long, porrect or obliquely ascending ; 
second joint thickened with appressed scales; terminal joint 
short, obtuse. Thorax with a small posterior crest. Abdomen 
with a series of small dorsal crests. Posterior tibiae with 
terminal spurs only. Forewings with 2 from #, 3 from near 
angle, 4 and 5 long-stalked from angle, 6 from upper angle, 
areole absent, 7, 8, 9, 11 stalked from before angle, 10 absent, 
11 running into 12. Hindwings with 2 from #, 3 and 4 
separate but approximated at origin, 5 from middle of cell, 
6 and 7 stalked, 8 anastomising with cell to ¢. 

Certainly one of the Gymmnoscelis group, and probably 
directly connected with Symmimetis, but in all other 


234 


Geometrites vein 5 of forewings arises from the middle, or 
above the middle of cell, with the exception of Microdes, 
in which it arises from below the middle, apparently in conse- 
quence of the development of some secondary sexual characters 
in the male. The stalking of 4 and 5 is an extraordinary 
anomaly in this family; possibly the discovery of the male 
may suggest some explanation. 


ANTIMIMISTIS ILLAUDATA, Nn. Sp. 

slaudatus, obscure. 

Q, 20-22 mm. Head grey. Palpi 14; whitish-ochreous 
sometimes greenish tinged. Antennae grey. Thorax grey. 
Abdomen grey; dorsum of second segment pale greenish- 
ochreous. Legs grey; anterior pair fuscous with whitish 
annulations on tarsi. Forewings triangular, costa nearly 
straight, gently arched towards apex, apex rounded, termen 
bowed, oblique; fuscous-grey with obscure whitish lines; first 
from 4 costa to 4 dorsum, indistinct, wavy; second from 
2 costa to % dorsum, slender, outwardly bowed, irregularly 
dentate; a fine parallel fuscous line succeeds this, and then a 
pale suffused line; a fine dentate subterminal line; cilia 
fuscous-grey. Hindwings with termen rounded, wavy; as 
forewings. Underside similar but more suffused. 

North Queensland: Kuranda, near Cairns, in November 
and April; two specimens received from Mr. F. P. Dodd. 


SYMMIMETIS MUSCOSA, Turn. 


North Queensland: Kuranda, near Cairns, in October ; 
Evelyn Scrub, near Herberton, in December. Queensland: 
Brisbane, in April. 


SYMMIMETIS SYLVATICA, N. sp. 
sylvaticus, of the woods. 


3, 9, 18-21 mm. Head fuscous. Palpi fuscous, towards 
base ochreous-whitish. Antennae fuscous; ciliations in 
male 24. Thorax grey mixed with fuscous. Abdomen pale 
greenish-ochreous with some fuscous scales; tuft in male 
whitish. Legs whitish-ochreous; anterior pair fuscous with 
whitish-ochreous annulations on tibiae and tarsi. Forewings 
broadly triangular, costa gently arched, apex rounded, termen 
bowed, oblique; whitish-ochreous suffused with fuscous, which 
forms indistinct markings; a large fuscous basal patch; a 
dark-fuscous discal dot at 4 on end of cell, and near posterior 
edge of basal patch; immediately following this a broad, 
dentate, transverse, whitish-ochreous line, indistinct towards 
dorsum; a broad median fuscous fascia containing some 


® 


235 


blackish scales on yon defied posteriorly by a fine, whitish, 
crenate line from 3 costa to 3 ? dorsum, bent outwards in disc ; 

a fine fuscous parallel line follows this, then a suffused whitish- 
ochreous fascia; a fuscous terminal band containing a fine, 
dentate, whitish subterminal line; a terminal series of whitish- 
ochreous dots on veins; cilia pale fuscous barred with whitish- 


ochreous opposite veins. Hindwings with termen rounded, 


slightly wavy; pale greenish-ochreous with patchy brownish 
irroration and a few blackish scales; a blackish discal dot 
at 4; cilia whitish-ochreous. Underside whitish with fuscous 
discal dots, subbasal, median, postmedian, and terminal fus- 
cous fasciae, postmedian of forewing angled outwards in 
middle. 

North Queensland: Evelyn Scrub, near Herberton, in 
December, January, and February; eight specimens received 


from Mr. F. P. Dodd. 


GYMNOSCELIS LOPHOPUS, Turn. 
Gymnoscelis homogona, Turn., is a synonym. 
North Queensland: Cairns, Herberton, Townsville. 


Queensland: Brisbane. Not uncommon in the last locality. 
New South Wales: Lismore. 


é 
GYMNOSCELIS SUBRUFATA, Warr. 


Forewings with 11 free. | 
Queensland: Duaringa, Brisbane, in February; one 
specimen taken at rest on a gate. 


GYMNOSCELIS TANAOPTILA, Turn. 


I have received a female example from Kuranda in 
November like male but smaller (18 mm.); posterior tibiae 
with terminal spurs only. 


GYMNOSCELIS ACIDNA, Turn. 


Forewings with 11 running into 12. 
North Queensland: Cairns, Townsville. 


GYMNOSCELIS SPODIAS, N. sp. 

s7rodos, ashes. } 

‘3, 9, 13-16 mm. Head whitish; sides of face and palpi 
dark fuscous. Antennae grey, towards base whitish; cilia- 
tions in male 4. Thorax and abdomen grey-whitish. Legs 
whitish; anterior pair mostly fuscous with whitish tarsal 
annulations. Forewings triangular, costa gently arched, apex 
rounded, termen bowed, oblique; 11 anastomising with 12; 
whitish with grey-whitish suffusion and obscure markings; 


236 


very faintly marked whitish transverse lines, subbasal, ante- 
median outwardly bowed, postmedian outwardly bowed, 
double, subterminal sometimes dentate; a few scattered 
blackish scales; blackish spots on costa near base, 1, 2, middle, 
and %, that on middle larger; a blackish spot in dise beneath 
second costal spot following subbasal line; a large blackish 
spot beneath mid-costa preceding postmedian line; cilia grey- 
whitish. Hindwings obtusely incised on vein 5, and with a 
rounded prominence on vein 4; as forewings but with one 
blackish spot preceding postmedian line, which forms a 
rounded projection in middle. Underside whitish partly 
suffused with grey. | 

Near G. acidmas, but much paler, lines much more 
obscure, except where partly defined by blackish spots. 

North Queensland: Evelyn Scrub, near Herberton, in 
December; Atherton. Queensland: Montville (1,500 ft.), 
near Nambour, in March. New South Wales: Stanwell Park, 
in April (Lyell). Four specimens. 


GYMNOSCELIS KENNiI, n. sp. 

@, 16 mm. Head brown; face and palpi blackish. 
Antennae pale brown. Thorax brown. Abdomen brown, 
dorsum suffused with blackish except towards base; tuft 
brown. Legs pale brown. Forewings triangular, costa nearly 
straight, towards apex arched, apex rounded, termen slightly © 
bowed, crenulate, strongly oblique; 11 running into 12; pale 
brown; markings and a few scattered scales blackish; a costal 
streak from base to beyond middle; a line from 4 costa, bent 
inwards beneath costa, thence strongly oblique to near base 
of dorsum; a second line from 3% costa, at first outwardly 
oblique, strongly bent inwards on vein 6, forming a second 
prominence on vein 4, bent outwards a third time above 
dorsum, ending on ? dorsum; a broad dark-fuscous suffusion 
from beneath costa beyond second line, broadening to fill whole 
tornal area; cilia brownish barred with blackish on crenula- 
tions. Hindwings with termen slightly rounded, wavy; pale 
brown densely suffused with dark fuscous beyond second line ; 
three blackish transverse lines, first subbasal, second at 4, 
third at 2 bent outwards beneath costa and again in middle; 
cilia brownish mixed with dark fuscous. Underside brownish 
suffused with fuscous without distinct markings. 

Exceptionally distinct. The broad. dark-fuscous suffusion 
of hindwings at once distinguishes it. 

Queensland: Gayndah, in October; one specimen received 
from Dr. Hamilton Kenny, an ardent naturalist and a per- 
sonal friend, to whom I dedicate it. . 


237 


GYMNOSCELIS HOLOCAPNA, 0. sp. 
6Aoxamrvos, Wholly smoky. 


g, 17-18 mm. Head fuscous. Palpi scarcely over 1; 
dark fuscous mixed with whitish-ochreous. Antennae grey; 
ciliations in male minute. Thorax and: abdomen fuscous- 
brown. Legs whitish-ochreous; anterior pair fuscous 
anteriorly. Forewings rather narrowly triangular, costa 
gently arched, apex rounded; termen bowed, oblique; 11 
running into 12; fuscous-brown or pale fuscous, markings 
obscurely darker; a basal patch; a moderate fascia at }, 
angled inwards beneath costa; a line from 3 costa, at first 
outwardly bowed, then slightly sinuate to #3 dorsum; a very 
obscure pale dentate subterminal line preceded by darker 
shading; cilia with basal half fuscous barred with whitish- 
ochreous opposite veins, terminal half grey. Hindwings rather 
narrow, termen strongly and evenly rounded; colour and 
cilia as forewings, but markings even more obscure; post- 
median line with a median tooth, indented below middle; 
subterminal line strongly dentate; some blackish irroration 
on dorsum. Underside fuscous-whitish. 


An obscure species. 
Northern Territory: Darwin, in September, December, 
and March; four specimens received from Mr. F. P. Dodd. 


CHLOROCLYSTIS PHOENOCHYTA, N. sp. 
gowoxvtos, suffused with reddish. 


@, 15 mm. Head whitish; face pale red. Palpi 2; 
grey. Antennae with joints expanded at apices; grey. 
Thorax whitish with a fine, transverse, postmedian line of 
dark-fuscous and reddish scales. [Abdomen and legs broken 
off.| Forewings elongate-triangular, costa slightly arched, 
apex round-pointed, termen bowed, oblique; 11 running into 
12; whitish partly suffused with grey and reddish ; costal edge 
reddish with some whitish strigulae; a broad, subbasal, grey 
fascia; its anterior edge outwardly curved, irregular; its 
posterior edge from 4 costa to 4 dorsum, forming a rather 
large posterior tooth beneath costa, beneath this obtusely 
indented; median area paler with indications of a suffused 
grey median line; a grey line from #% costa to % dorsum, 
strongly outwardly curved, slightly dentate; this is followed 
by a fine, parallel, dentate, grey line; disc beyond this 
suffused with pale red; a whitish, dentate, subterminal line; 
an interrupted grey terminal line; cilia pale reddish mixed 
with grey, apices grey-whitish. Hindwings with termen 
rounded ; wholly suffused with pale red except extreme base ; 
some few dark-fuscous scales on veins; a pale transverse line 


238 


at 4; another, broader, at 3 containing a very fine reddish 
line; subterminal indistinct, ut preceded by grey dentations; 
cilia pale reddish, apices grey-whitish. 

This species is very distinct by the ‘red suffusion, but, 
the posterior legs being absent, 1t is not possible to be sure 
that it is not a Gymnoscelis, Type in Coll. Lyell. 

Northern Queensland:, Gordonvale, near Cairns; one 
specimen. . 

CHLOROCLYSTIS EURYLOPHA, Nl. sp. 

etpvAodos, broadly crested. 

3,9, 15-16 mm. Head pale grey. Palpi 21; pale grey 
with a few darker scales. Antennae whitish-grey. Thorax 
and abdomen grey. Legs ochreous-whitish; anterior pair 
mostly grey; outer median spur 4. Forewings triangular, 
costa rather strongly arched, apex round-pointed, termen 
bowed, oblique; pale grey with numerous, wavy, fuscous, 
transverse lines more or less distinct; costa of male with a 
crest of long hairs extending from near base to middle; trans- 
verse lines in basal half of wing sometimes very indistinct, 
but sometimes as many as six can be distinguished, all out- 
wardly curved; a more distinct line from 2 costa, at first 
outwardly oblique, forming two short, obtuse, posterior pro- 
jections; then inwardly oblique to 2% dorsum; several paler 
indistinct lines follow this; an obscure, pale, dentate, sub- 
terminal line; a fuscous terminal line, interrupted on veins; 
cilia pale grey. Hindwings with termen scarcely rounded, 
irregularly waved; as forewings. 

This little species requires careful discrimination. The 
male may be distinguished readily from C. epilopha by the 
much wider extent of the crest on costal margin of forewing. 
Between the female of these two species it is hard to give any 
distinction, but the presence of blackish scales on the veins 
in the basal part of forewing in epilopha is helpful. The 
female also somewhat resembles C. ansigillata, but the 
rounded and not waved termen of the hindwing in the latter 
is in itself sufficient ‘difference. 

Queensland : Montville, near Nambour, in March; seven 
specimens (one male, six females). 


CHLOROCLYSTIS PYRSODONTA, Nl. Sp. 
mupoodovtos, with reddish tooth. 


3,9, 15-16 mm. Head fuscous. Palpi 14; whitish- 
ochreous mixed with blackish towards base. Antennae grey ; 
ciliations in male minute. Thorax pale grey, anterior edge 
fuscous. Abdomen pale grey. Legs fuscous; posterior pair 
paler; outer spurs about 4 of inner spurs. Forewings broadly 


239 


triangular, costa gently arched, apex rounded, termen bowed, 
oblique; whitish, markings extremely pale grey, except in 
costal +, where they are fuscous and distinct; a fuscous costal 
streak from base to first fascia; first fascia at 4, moderately 
broad, sharply angled inwards beneath costa; second fascia 
median, similar to first, lke it sharply angled inwards 
beneath costa; third fascia beyond #, narrower except on 
costa, evenly curved, posteriorly limited by a finely dentate, 
whitish, subterminal line; a fine fuscous terminal line inter- 
rupted on veins; cilia grey, apices paler. Hindwings with 
termen unevenly rounded; concave above middle, prominent 
between veins 3 and 4; as forewings; but median fascia 
reddish with a few blackish scales, and a strong, obtuse, 
median, posterior tooth; without dark costal markings. 
Underside pale fuscous, with a darker, posteriorly toothed, 
median, transverse fascia on hindwings. 


Northern Queensland: Cardwell, one wasted hs in 
August; Evelyn Scrub, near Herberton, male type, in 
January (F. P. Dodd). 


CHLOROCLYSTIS NIGRILINEATA, Warr. 


36,92, 18 mm. Head whitish-grey. Palpi about 1; 
whitish-grey mixed with blackish. Antennae whitish-grey. 
Thorax whitish-grey. Abdomen whitish-grey with some in- 
constant dark-fuscous markings. Legs ochreous-whitish ; 
anterior pair grey. Forewings triangular, moderately broad, 
costa slightly arched, apex round-pointed, termen bowed, 
oblique; 11 running into 12; whitish-grey with pale-grey and 
dark-fuscous transverse lines; a dark-fuscous subbasal line 
with median posterior tooth; a dark-fuscous wavy line from 
4 costa to + dorsum; a pale-grey median line, sometimes 
double; a dark-ftscous line from costa before %, with two 
obtuse posterior teeth, subcostal and median, thence oblique 
and slightly dentate to # dorsum; a very faint, pale, dentate 
subterminal line preceded by an interrupted dark-fuscous 
line; a terminal series of interneural fuscous dots; cilia pale 
grey. Hindwings with termen rounded; as forewings but all 
lines indistinct except postmedian, which has a posterior 
angular projection. about middle. Underside pale grey, 
darker towards termen, with fuscous postmedian lines on both 
wings. 

_ My examples agree well with Warren’s description. The 
dark transverse lines are conspicuous. 

Northern Territory: Darwin, in November and February ; 
two specimens received from Mr. F. P. Dodd. Queensland: 
Duaringa (Warren). 


240 


CHLOROCLYSTIS POLIOPHRICA, Nl. Sp. 
todopptkos, grey-rippled. 


3, @, 13-16 mm. Head whitish. Palpi whitish, in 
male annulated, in female irrorated with dark fuscous. 
Antennae whitish, towards apex tinged with grey; ciliations 
in male minute. Thorax pale fuscous; patagia whitish. 
Abdomen whitish with some fuscous scales. Legs whitish ; 
anterior pair fuscous; posterior tibiae with inner spurs long, 
outer spurs 4, outer median spur absent in male. Forewings 
in male with costa straight in basal half, strongly arched in 
apical half, in female evenly arched throughout, apex 
rounded, termen bowed, oblique; whitish with fuscous mark- 
ings; basal # of costa more or less suffused; a number of 
indistinct transverse lines preceding postmedian, in male 
obsolete towards. dorsum; postmedian line from % costa, at 
first outwardly oblique, forming two angular posterior pro- 
jections in disc, thence inwardly oblique to % dorsum; a 
fuscous subterminal line, in male thickened into spots beneath 
costa, above middle, and below middle, interrupted between 
spots, in female more uniform; an interrupted terminal line; 
cilia whitish, in male with some obscure fuscous bars. Hind- 
wings with termen gently rounded, slightly wavy; as fore- 
wings; postmedian line with an angular indentation above 
middle, and an angular projection in middle. Underside 
fuscous-whitish. 

Queensland: Dulong, near Nambour, in December, one 
female; Brisbane, in April, one male type. 


Gen. Micropes, Gn. 


This genus has two remarkable peculiarities in the neura- 
tion of the forewing. One is the approximation of vein 5 
at its origin to 4. This is probably secondary to the peculiar 
sexual modification in the forewing of the male. The other is 
that 11 runs into 12 in villosata and asystata, but has second- 
arily disappeared altogether in squamulata, diplodonta, and 
ortochares,; typhopha and melanocausta T have not examined. 


MICRODES ORIOCHARES, nN. sp. 

6pecoxapys, rejoicing in the mountains. 

3, 9, 18-20 mm. Head dark fuscous. Palpi in male 4, 
in female 44; dark fuscous. Antennae fuscous; in male 
thickened and slightly laminate, ciliations }. Legs fuscous; 
anterior pair dark fuscous; anterior and middle tarsi with 
ochreous-whitish annulations. Forewings with costa moder- 
ately: and evenly arched, apex round-pointed, termen bowed, 
moderately oblique; fuscous; a slender, obscure, outwardly 


241 


curved, transverse line at } followed by a pale, indistinctly 
double line; beyond this isa brownish-tinged fascia, not 
always developed ; beyond this a paler area containing two or 
three very obscure, slender, transverse lines; a whitish line 
edged posteriorly by a dark fuscous line from # costa, at first 
moderately outwardly oblique, acutely angled outwards above 
middle, thence concave to below middle, where it is again 
angled outwards, thence straight to ~ dorsum; a slight 
brownish suffusion on posterior edge of this line; a fine, 
irregularly dentate, whitish, subterminal line; cilia fuscous, 
sometimes very obscurely barred, apices grey. Hindwings 
with termen strongly ae slightly wavy; pale grey; an 
obscure darker line at ¢; cilia pale grey. 

Certainly near J. apie ania: Turn., but smaller, fore- 
wings’ proportionately broader, less brownish, costa less 
strongly arched, cilia not distinctly barred, palpi in male 
rather longer. Unless intermediate forms are discovered it 
should be regarded as a distinct species. 

New South Wales: Mount Kosciusko, in January, 
February, and March; seven specimens. Victoria: Mount 
St. Bernard (5,000 ft.), in February ; a large female (24 mm.) 
in Coll. Lyell. 

MICRODES ASYSTATA, N. Sp. 


aovoratros, inconstant. 


2, 26-30 mm. Head, thorax, and abdomen fuscous with 
scanty whitish irroration. Palpi ‘3k; second joint expanded 
by rough scales above and beneath; terminal joint short; 
fuscous irrorated with whitish. Antennae fuscous. Legs 
fuscous; tarsi with fine whitish annulations; posterior pair 
ochreous-whitish. Forewings triangular, costa gently arched, 
apex round-pointed, termen straight, very slightly oblique; 
whitish irrorated with grey; numerous fine transverse 


_fuscous lines more or less distinct; sometimes stronger lines 


define median area; first from 4 costa to 4 dorsum, outwardly 
curved; second from 4 costa to tornus, with a small acute 
posterior tooth beneath costa, and an obtuse tooth beneath 
middle; sometimes median area is partly or wholly fuscous,. 
and lines indistinct; a finely dentate, whitish, subterminal 
line; cilia grey. Hindwings with termen strongly but un- 
evenly rounded, projecting slightly on veins 3 and 6; grey; 
cilia grey. 

Male unknown and female inconstant; in one example 
the anterior margin of median band is much more strongly 


rounded posteriorly, an unusual form of variation. Type in 


Coll. Goldfinch. f | 
New South Wales: Mount Kosciusko, in February; three 
specimens. 


| 242 


Gen. Scorocyma, Turn. 

This comes near the European genus Hyirrhoé, Hb., but 
differs in 7, 8, 9, 10, and 11 arising by a common stalk from 
the small areole. 

ScOTOCYMA ALBINOTATA, WIk. 

Mr. Prout informs me that Paragramma mimula, Warr., 
is a synonym. 

SCOTOCYMA IDIOSCHEMA, Ni. 6p. 

idvorxnuos, of peculiar pattern. 

Q, 31-34 mm. Head whitish-brown mixed with dark 
brown. Palpi slightly over 1; whitish-brown irrorated with 
dark fuscous. Antennae grey. Thorax brown; patagia partly 
whitish-brown. Abdomen brown. Legs whitish-ochreous ; 
anterior pair fuscous with whitish-ochreous basal annulations. 
Forewings triangular, costa gently arched, apex rounded, 
termen bowed, slightly oblique, crenulate; a fuscous basal 
patch to +, containing some whitish-ochreous transverse lines 
on costa prolonged to middle, and with an inferior tooth near 
extremity; remainder of disc except a costal strip, and tri- 
angular apical and tornal areas occupied by a very large 
whitish-ochreous blotch, suffused with brown, or dark 
ferruginous-brown except at edges; costal strip fuscous 
strigulated with whitish-ochreous; dorsal edge narrowly and 
interruptedly fuscous; apical and tornal triangles fuscous- 
brown, containing an incomplete, fine, dentate, ochreous- 
whitish line, sometimes forming a white spot above tornus, 
a white spot sometimes present on margin of central blotch 
above tornus; cilia fuscous partly mixed with whitish- 
ochreous. Hindwings with termen rounded, dentate; 
brownish; some whitish dots on veins; sometimes obscure 
pale-fuscous transverse lines; some variable white spots pre- 
ceding termen; a dark-fuscous terminal line; cilia fuscous- 
brown. Underside whitish with many, more or less distinct, 
transverse lines and a broad subterminal fascia fuscous. 

North Queensland: Kuranda, in November (Coll. Lyell) ; 
Evelyn Scrub, near Herberton, in October. Queensland: 
Brisbane, in January. Three specimens. 


ScoTOCYMA EURYOCHRA, Nn. sp. 

evpuwxpos, broadly pale. 

Q, 34 mm. Head dark fuscous. Palpi 1; dark fuscous 
with a few whitish scales. Antennae fuscous. Thorax fus- 
cous. Abdomen grey; apex fuscous. Legs fuscous; tarsi 
with fine ochreous-whitish annulations; posterior pair mostly 
ochreous-whitish. Forewings triangular, costa moderately 
arched, apex rounded-rectangular, termen bowed, slightly 


243 


oblique, crenulate; grey-whitish with numerous, fine, indis- 
tinct, wavy, transverse lines; markings brownish-fuscous; a 
rather large basal patch containing some grey- “hee suffusion , 
limited by a slightly eurved wavy line from 4 costa to 4 dor- 
sum; median band ill-defined, mostly grey- -whitish with fine 
lines, but with some fuscous suffusion on costa ; a large apical 
blotch; two subterminal spots above tornus; cilia fuscous, 
towards centre of termen partly grey-whitish. Hindwings 
with termen rounded, slightly dentate; as forewings; basal 
patch very small; a broad terminal band, containing a sub- 
terminal series of whitish dots con veins; a dark-fuscous 
terminal line, interrupted by whitish dots on veins; cilia 
brownish-fuscous. 

New South Wales: Toronto, near Newcastle, in April; 
one specimen. Type in Coll. Goldfinch. 


Gen. CHAETOLOPHA, Warr. 


Type C. oxyntis, Meyr. This name must be adopted for 
the small endemic genus, to which I formerly applied the 
name Scordylia, Gn. The areole is large and 11 widely 
separate. In Hulype, Hb., type hastata, Lin., which other- 
wise resembles it in neuration, the areole is smaller and 11 
near or connate from its apex. There is, I think, no really 
close relationship between the two genera. The species of 
Chaetolopha are narrow-winged; in the males of oxyntis and 
leucophragma there is a small subterminal scale-tuft on vein 2 
of hindwings on underside, but this is absent in niphosticha 
and emporias; of the other two species I have no male to 
examine. The penultimate abdominal segment of the male 
bears a pair of lateral tufts. By boiling in potash- the 
abdomen of the male lewcophragma is shown to bear a pair 
of extrusible scent-organs on the fourth segment. In the 
male of niphosticha the termen of the hindwings is produced 
to form an acute central tooth. 


Gen. Eccymatoce, Prout. 


Prout, Ann. Transvaal Mus., ii., p. 207 (1913). 
Type #. melanoterma, Prout, from South Africa. 


EcCCYMATOGE CALLIZONA, Low. 


I am now satisfied that the type of fulvida, Turn., is 
merely an aberration of callizona. 


ECCYMATOGE MORPHNA, D. sp. 
popdvos, dusky. 
36,30 mm. Head fuscous; face dark fuscous with a few 
whitish scales. Palpi 11; dark fuscous with a few whitish 


4 


244 


scales. Antennae fuscous; in male thickened and minutely 
ciliated. Thorax and abdomen fuscous; anal valves in male 
large. Anterior legs dark fuscous [middle and posterior pairs 
broken off]. Forewings triangular, costa gently arched, apex 
acute, termen bowed, oblique, finely dentate; fuscous; mark- 
ings dark fuscous, obscure; a small basal patch; a median 
fascia containing a darker discal dot beneath mid-costa, 
limited anteriorly by a nearly straight line from 4 costa to 
4 dorsum, posteriorly by a line from 2 costa, strongly bent 
outwards in disc, with an angular prominence between veins 
7 and 8, and another between 3 and 4, thence sinuate to 
# dorsum; an indistinct pale subterminal line; a terminal 
line interrupted by pale dots on veins; cilia fuscous. Hind- 
wings with termen slightly rounded, irregularly dentate, with 
a stronger tooth on vein 4; as forewings, but without discal 
dot, and posterior edge of median fascia only slightly un- 
dulating. Underside fuscous without markings. 

A true Ececymatoge ; though vein 5 of hindwings is scarcely 
from below the middle of cell, the upper discocellular is bent. 
There appears to be no thoracic crest. 


New South Wales: Mount Kosciusko (3,500 ft.), in 
January; one specimen. 


Gen. Horisme, Hb. 
Hucymatoge, Sect. I., Turn., Proc. Roy. Soc. Vict., 
1903, p. 247. 
Differs from Hucymatoge in the presence of a posterior 
thoracic: crest. 
HoRISME MORTUATA, Gn. 


3, Q, 27-30 mm. Very similar to H. scotodes, Turn., 
but slightly larger; palpi longer, 24 to 3 as against 14 to 2 
in the latter; forewings with basal lines more evenly curved, 
in the latter they are more oblique; postmedian line with a 
doubly obtuse-toothed projection. 

New South Wales: Sydney, in January and February. 
Victoria: Beaconsfield. Three examples. 


HORISME PLAGIOGRAPHA, Nl. sp. 
tAaytoypados, obliquely inscribed. 


Q, 25-26 mm. Head grey. Palpi 3; grey irrorated 
with dark fuscous, whitish beneath towards base. Antennae. 
grey. Thorax grey mixed with fuscous, posterior edge of 
crest fuscous. Abdomen grey, some fuscous scales in crests. 
Legs whitish, on dorsal surfaces fuscous. Forewings tri- 
angular, costa straight, slightly sinuate before apex, apex 


245 


acute, termen bowed, oblique, slightly crenulate; whitish 
with fuscous suffusion and markings, a conspicuous dark- 
fuscous oblique bar from dorsum near base to middle of disc, 
sometimes forming a complete fascia to 4 costa; two or three 
fine, incomplete, transverse lines between this and costa; a 
nearly straight band of three fine fuscous lines from mid-costa 
to dorsum at +; a dark-fuscous, median, subcostal, discal dot ; 
a suffused band, towards costa resolvable into three lines, 
from # costa to 2 dorsum, outwardly curved with slight obtuse 
prominences above and below middle; a streak from apex to 
upper prominence on postmedian line, slightly downwardly 
curved; between this,and costa is a paler apical area; sub- 
terminal whitish, very ill-defined; a terminal line; cilia grey- 
whitish with a few darker points. Hindwings with termen 
very little rounded, dentate; whitish-grey, with fine, fuscous, 
transverse lines from dorsum, becoming indistinct before 
costa; from' 4, middle, an outwardly-curved, stronger line 
from %, a fine line following close on this, and a double 
subterminal line; a fuscous terminal line; cilia whitish-grey. 


New South Wales: Sydney (Manly), in October; Jervis 
Bay; in September; two specimens. There is a third female 
example from the latter locality in Coll. Goldfinch, taken in 
November. Type in Coll. Lyell. 


Gen. Crparia, Treit., Kur. Schmet., vi., 2, p. 140. 


Mr. L. B. Prout informs me that the type is the Euro- 
pean C. fulvata, Forst., and that Hydriomena, Hb., is a 
synonym of later publication. This large European genus is 
but poorly represented in Australia, and in New Zealand 
there is no endemic species, the only representative there 
being subochraria, Dbld. Six Australian species are known ; 
of these swbhochraria, apotoma, uncinata, and microcyma are 
probably derived through the Antarctic; scythropa and /asio- 
placa are not nearly allied specifically to the first four, and 
entered Australia from the north. The groove on hindwing 
of male of scythropa I consider a character of specific value 
only. Heterochasta, Meyr., and Polyclysta, Gn., are deriva- 
tions of this second section of the genus. 


LARENTIA PETRODES, Turn. 


Queensland: Warwick. Victoria: Gisborne. 


LARENTIA XERODES, Meyr. 


I have examined what I believe to be an example of 
Xanthorhoé xerodes, Meyr., and refer it to this genus. 


246 


LARENTIA ORIBATES, N. sp. 

opeBaryns, a mountaineer. 

6, 28 mm. Head whitish irrorated with fuscous; face 
blackish. Palpi 14; dark fuscous. Antennae grey; pectina- 
tions in male 6, extreme apex simple. Thorax dark fuscous 
irrorated with whitish. Abdomen grey-whitish; paired fus- 
cous dots on the dorsum of each segment except the first two. 
Legs grey-whitish; anterior and middle pairs fuscous on 
dorsum. Forewings triangular, costa nearly straight, slightly 
arched towards apex, apex pointed, termen longer than 
dorsum, slightly bowed, slightly oblique; whitish with 
numerous fine, fuscous, oblique, transverse lines; costa irror- 
ated with fuscous; a line from } costa to near base of dorsum; 
another parallel from 2 costa; a median band of three or four 
close lines, anterior edge from mid-costa to + dorsum, nearly 
straight, posterior from # costa to before middle of dorsum, 
slightly curved outwards in middle of disc; beyond and parallel 
is a very fine line thickened by some small dots; beyond this 
again three close, parallel, wavy lines; an oblique fuscous 
shade from apex; a narrow grey terminal fascia; an inter- 
rupted fuscous terminal line; cilia whitish with a grey median 
line. Hindwings with termen rounded; as forewings but all 
lines, except terminal line, becoming obsolete in costal area, 
which is whitish, and in male contains an oval patch of 
ochreous-grey altered scales. | 

Victoria: Mount St. Bernard, in February; one specimen 
received from Dr. W. E. Drake. 


LARENTIA AGANOPIS, 0. sp. 
ayavwms, gentle-looking. 


3,9, 24-32 mm. Head whitish. Palpi in male l, 
in female 11; grey-whitish. Antennae pale grey; pectina- 
tions in male 5, extreme apex simple. Thorax ochreous- 
whitish. Abdomen ochreous-whitish, suffused with pale grey 
on dorsum. Legs fuscous; tarsi annulated with whitish; 
posterior pair whitish. Forewings triangular, costa gently 
arched, apex round-pointed, termen bowed, oblique; ochreous- 
whitish; markings pale grey, brownish tinged; a very small 
basal patch, followed by two fine parallel lines confluent on 
costa; median band broad from costa to middle, much nar- 
rower from middle to dorsum, darker on costa; anterior edge 
from 4 costa to + dorsum, outwardly curved; posterior edge 
from 2% costa, obtusely toothed beneath costa, with a slight 
double-toothed median prominence, thence strongly oblique 
and dentate to mid-dorsum; this is followed by two fine in- 
distinct parallel lines; an indistinct pale subterminal line 


al 


247 


preceded by a slight dark suffusion towards costa; a terminal 
series of triangular marks or fine, short, interneural, longi- 
tudinal streaks; cilia ochreous-whitish. Hindwings with 
termen rounded; ochreous-whitish, with pale, suffused, 
median, postmedian, and submarginal grey lines; terminal 
marks and cilia as forewings. Underside whitish; forewings 
suffused with grey as far as postmedian line; hindwings with 
a median transverse line. 

New South Wales: Woodford, in March and April; two 
specimens received from Mr. Geo. Lyell. Type in Coll. Lyell. 


Gen. Meuitutias, Meyr. 

I do not consider the presence of androconial scales in 
the male as a rule a sufficient character for generic distinction, 
and have therefore merged Hypycnopa, Low., in Xanthorhoé, 
and refrained from making a new genus for Larentia petrodes. 
But I have sacrificed strict consistency in retaining the genus 
Melitulias, Meyr., which defines a small natural group peculiar 
to Tasmania and South-east Australia, particularly the 
mountains, in which new species may be expected to occur. 
It is an endemic derivative of Huphyia. I regard glandulata, 
Gn., as the type. 


MELITULIAS LEUCOGRAPHA, 0. sp. 

Aevkoypados, inscribed with white. 

3, 9, 24-28 mm. Head fuscous with a few whitish scales 
on face. Palpi 3; fuscous; base beneath whitish, sharply 
defined. Antennae dark fuscous; ciliations in male imper- 
ceptible. Thorax and abdomen fuscous with a few whitish 
scales. Legs fuscous. Forewings triangular, costa very 
slightly arched, apex pointed, termen bowed, oblique, wavy ; 
fuscous-brown; markings white partly outlined with fuscous ; 
a fine line from + costa to + dorsum, curved outwards beneath 
costa; a broader line from 4 costa, at first transverse, then 
bent inwards and joining first line above dorsum; a dark- 
fuscous discal dot beneath mid-costa, sometimes surrounded by 
a narrow whitish suffusion ; sometimes a fine, sinuate, inwardly 
oblique line from 2 costa not reaching middle of disc; a 
broader line from 2 costa, angled inwards above and outwards 
at middle, then inwardly curved to dorsum before tornus; a 
fine interrupted subterminal line, a fine oblique streak from 
apex, crossing subterminal line, ending in postmedian line at 
its subcostal angle; a dark-fuscous terminal line; cilia fuscous 
barred with white. Hindwings with termen rounded; in 
male grey; with a large, median, oval, brownish-fuscous, 
androconial blotch; cilia grey; in female pale brownish-grey, 
a suffused, whitish, postmedian, transverse line; a whitish 


248 


subterminal line; terminal line and cilia as forewings. Under- 
side of hindwings in both sexes like upperside in female, but 
more distinctly marked and with a dark-fuscous antemedian 
discal dot. 

Near /. graphicata, Wik., but easily distinguished by 
the hindwings. 

New South Wales: Mount Kosciusko (5,000 ft.), in 
December; three specimens. Type in Coll. Goldfinch. 


Gen. Hupnyia, Hb., Verz., p. 326. 


Type /. mcata, Hb., from Europe. This genus corre- 
sponds to Hydriomena, Section I., of my revision. In Aus- 
tralia it is the dominant genus of the family, being especially 
~ well represented in South-east Australia and Tasmania; many 
more species will doubtless be discovered, especially in the 
mountains. The genus is also moderately well represented in 
New Zealand. 

EuPHyIA SymMpHONA, Meyr. 


Epirrhoé maerens, Swin. (Trans. Ent. Soc., 1902, p. 648), | 
is a synonym (teste Prout, in lit. ). 


EUPHYIA TACERA, Nn. sp. 

Taxepos, soft. 

3, 9, 30-32 mm. Head brownish; face fuscous. Palpi 
2; fuscous; beneath whitish-ochreous. Antennae fuscous; 
ciliations in male minute. Thorax and abdomen brownish- 
fuscous. Legs-fuscous; tarsi annulated with ochreous-whitish. 
Forewings triangular, costa moderately arched, apex round- 
pointed, termen bowed, slightly oblique; whitish partly 
suffused with pale brownish ; a small brown basal patch limited 
by a fine fuscous line; two ill-defined, very fine, transverse, 
fuscous lines follow this; median band rather ndrrow, brown 
with fine fuscous transverse lines, sometimes with a narrow 
central grey band; anteriorly limited by a fine, slightly 
outwardly-curved line from 4 costa to 4 dorsum, posteriorly 
by a similar line from before 2 costa to before 3 dorsum, with 
slight rounded prominence beneath costa, and again in 
middle; this is followed by a suffused whitish band containing 
two suffused, wavy, fuscous, transverse lines; a broad brownish 
terminal suffusion, containing a finely crenulate, whitish, sub- 
terminal line, preceded and followed by slight fuscous suf- 
fusion ; a fuscous oblique mark beneath apex; cilia brownish- 
grey, apices pale grey. Hindwings with termen rounded, 
wavy; yellow-ochreous; three fine fuscous transverse lines 
from basal half of dorsum, of which only the first reaches 
costa; a double subterminal line from dorsum usually reaching 


es 


RSI 


249 


about middle; a narrow terminal band, sometimes. obsolete 
towards apex; a dark-fuscous terminal line obsolete towards 
apex ; cilia fuscous, towards apex pale yellow. 

Not unlike #. lucidulata, Wlk., which may be at once 
distinguished by the indented antemedian line. | 

New South Wales: Barrington Top, in December ; three 
specimens. Type in Coll. Goldfinch. 


EUPHYIA PERIALLA, 0. sp. 
mepiadAos, excelling. 


3, 9, 30-35 mm. Head fuscous. Palpi 24; fuscous, at 
base whitish beneath. Antennae fuscous; ciliations in male 
minute. Thorax fuscous. Abdomen fuscous, beneath 
ochreous-whitish. Legs fuscous irrorated, and tarsi annulated 
with whitish-ochreous. Forewings broadly triangular, costa 
moderately arched, apex round-pointed,: termen bowed, 
oblique, wavy; brown with fuscous and whitish lines; a small 
basal patch defined by a transverse, outwardly curved line; a 
slightly paler fascia follows this; median band fuscous, 
broad on costa but narrow on dorsum, containing a paler 
costal area defined by a fuscous line extending nearly to 
middle, with a blackish discal mark near its anterior edge; 
fine whitish lines defining median band, anterior from 3 costa 
to 4 dorsum, outwardly curved, posterior from beyond 4% costa 
to before dorsum, at first transverse, then shortly incurved, 
and forming an obtuse double prominence in middle; two 
fine parallel fuscous lines follow this; a fine, interrupted, 
_ whitish, subterminal line, preceded and near apex followed 
by some fuscous suffusion; a dark-fuscous terminal line in- 
terrupted on veins; cilia fuscous with a whitish basal line, 
apices with obscure pale bars. Hindwings with termen 
strongly rounded, dentate; orange; towards dorsum suffused 
with fuscous containing many darker and paler short trans- 
verse lines; this suffusion extends on termen to middle; ter- . 
minal line and cilia as forewings, but paler towards apex. 
Underside pale ochreous partly suffused with fuscous; both 
wings with discal dot, transverse lines, and terminal band 
fuscous, the last containing a slender; ‘whitish, subterminal 
line. 

New South Wales: Mount Kosciusko (4,500 ft.), in 
January; one male. Victoria: Mount St. Bernard, in 
February; two females, in Coll. Lyell. Two specimens from 
New South Wales (Ebor) in January and Victoria (Castle- 
maine, Dr. W. E. Drake) in March are probably of the 
same species, but the forewings are much paler except in | 
basal patch and median band. Two since received from Mr. 
H | 


a m 


250) 


G. W. Goldfinch taken on Barrington Top. in December 
resemble the Kosciusko type. 


EUPHYIA SYMMOLPA, N. sp. 


ocvppodros, in harmony. 


29, 32 mm. Head fuscous; frons rounded ; slightly pro- 
jecting; frontal tuft whitish. Palpi 3; whitish mixed with 
fuscous. Antennae fuscous. Thorax and abdomen fuscous 
with fine whitish irroration. Legs fuscous with fine whitish 
irroration; posterior pair mostly whitish. Forewings tri- 
angular, costa straight except close to base and apex, apex 
round-pointed, termen moderately bowed, moderately oblique, 
slightly undulating; pale fuscous with fuscous markings; a 
basal patch of three or four transverse lines; a short line from 
dorsum to cell follows this; median band limited anteriorly 
by a double, nearly straight line from 4 costa to mid-dorsum, 
posteriorly by a double line from beyond 4 costa, at first 
transverse, with a strong, angular, posterior projection in 
middle (in ene example there is a slighter angle also beneath 
costa), thence concave to # dorsum, this line is edged pos- 
teriorly by a well-marked whitish line; a blackish discal spot 
in median band beneath mid-costa; a strong, crenulate, 
whitish, subterminal line from costa shortly before apex to 
tornus, edged anteriorly by a series of fuscous spots; a dark- 
fuscous terminal line; cilia fuscous, apical % barred with 
whitish. Hindwings with termen slightly rounded, slightly 
undulating ; whitish, towards margins grey; a grey discal dot 
at 1; an ill-defined grey terminal band containing an un- © 
dulating whitish line; terminal line and cilia as forewings. 

Not unlike (. symphona, Meyr., but differing in the — 
form of postmedian line, discal spot not pale centred, and 
other details. ‘ 

New South Wales: Mount Kosciusko (6,000 to 7,000 ft.), 
in January; two specimens. 


EUPHYIA LEPTOPHRICA, Nl. Sp. 
Aerrodpixos, finely rippled. 


3, 9, 34-38 mm. Head, thorax, and abdomen grey. 
Palpi 24; dark grey, beneath whitish. Antennae grey; cilia- 
tions in male extremely short. Legs fuscous, irrorated, and 
tarsi annulated, with grey-whitish. Forewings broadly tri- 
angular, costa strongly arched, apex round-pointed, termen 
bowed, slightly oblique, wavy; grey, with numerous slender, 
finely crenulate, fuscous, transverse lines; basal patch hardly 
defined; median band obscurely defined, anteriorly by p 
slightly curved wavy line from 4 costa to 4 dorsum, posteriorly 


251 


by a similar line from #? costa to # dorsum, with a slight 
doubly subacute median projection ; a fuscous discal dot before 
middle; a fine, crenulate, whitish, subterminal line; a 
blackish terminal line, interrupted on veins; cilia grey. Hind- 


‘wings with termen rounded, wavy; pale grey with fine wavy 


transverse lines not reaching costa; terminal line and cilia as 
forewings . 
Type in Coll. Goldfinch. Perhaps nearest Z. NG Sti i 
Meyr. 
New South Wales: Pare Top, in December; two 
specimens. 
EKUPHYIA PANOCHRA, 1. sp. 


mavwxpos, Wholly pale. 


36, Q@, 28-32 mm. Head ochreous-whitish with a very 
few dark-fuscous scales. Palpi 23; ochreous-whitish with 
slight dark-fuscous irroration. Antennae ochreous-whitish 
annulated with fuscous; in male slightly thickened, cilia- 
tions 4. Thorax ochreous-whitish. Abdomen ochreous-whitish 
with a few pale-grey scales on dorsum. Legs ochreous-whitish 
irrorated with fuscous. Forewings broadly triangular, costa 
rather strongly arched, apex subrectangular, termen nearly 


a straight, slightly oblique; ochreous-whitish, with slight pale- 


grey suffusion, more distinct towards termen ; a very fine, often 
indistinct, slightly curved, slightly dentate, fuscous line from 
4 costa to 4 dorsum; a second, similar, nearly straight line, 
finely dentate, from # costa to 4 dorsum; in some examples a 
third line or series of fine dots beyond this; cilia ‘dark grey, 
apices white except on costa, beneath apex, and on tornus. 


- Hindwings with termen rounded; oehreous-whitish, without 


markings; cilia grey, apices whitish. Underside of forewings 
suffused with grey; of hindwings with grey oo discal 
dot, postmedian, and subterminal lines. 

New South Wales: Mount Kosciusko (5,000 ft.), in 
January. Victoria: Mount St. Bernard. (5,000 ft.), 
February; eight specimens. Type in Coll. Lyell. 


KUPHYIA OXYODONTA, D. sp. 
éévodovros, sharply-toothed. 
Q, 28 mm. Head pale grey. Palpi 2; whitish with 


- fuscous irroration. Antennae fuscous. Thorax whitish mixed 


with grey. Abdomen ochreous-whitish suffused with fuscous 

on dorsum. Legs fuscous irrorated, and tarsi annulated, 

with ochreous-whitish ; posterior pair mostly ochreous-whitish. 

Forewings triangular, costa gently arched, apex round-pointed, 

termen nearly straight, oblique, wavy ; whitish with fuscous 

markings; a small basal patch with three darker lines, one of 
H2 


252 


which forms its posterior edge, and is slightly rounded, 
slightly dentate, transverse; median band broad; its anterior 
edge broadly dark fuscous from + costa to 4+ dorsum, strongly 
concave, indented above and below middle; a linear ante- 
median discal mark followed by two fine incomplete fuscous 
lines; posterior edge marked by a fine dark-fuscous line, 
thickened above middle, from #? costa, projecting slightly 
beneath costa, then angularly indented, with a strong median 
double-toothed projection, the upper tooth more prominent 
and acute, thence inwardly curved and dentate to #? dorsum, 
suffused fuscous spots on costa before apex, in disc beneath 
this, on termen beneath apex, and above tornus; an inter- 
rupted terminal line; cilia whitish with a broad fuscous 
median line. Hindwings with termen slightly rounded, wavy; 
whitish-grey; four or five faintly darker transverse lines 
better marked on dorsum; postmedian line with a median 
acute tooth ; an interrupted fuscous terminal line; cilia whitish 
with some grey and fuscous scales. 

Western Australia: Perth, in April; one specimen 
received from Mr. W. B. Alexander. 


EUPHYIA POLIOPHASMA, N. sp. 
Todiopacpos, grey ghostly. 


3, 36-38 mm.; 9,32 mm. Head, thorax, and abdomen 
pale grey irrorated with fuscous. Palpi 24; fuscous, towards 
base ochreous-whitish. Antennae with internal surface fuscous, 
external whitish ; in male shortly laminate, ciliations }. Legs 
pale grey irrorated with fuscous. Forewings triangular, costa 
gently arched, apex round-pointed, termen bowed, slightly 
oblique; pale grey with slight fuscous irroration ; antemedian 
line obsolete; postmedian slender, fuscous, crenulate, slightly 
projecting in middle, from 2 costa to % dorsum, sometimes 
obsolete ; Gilia grey. Hindwings with termen rounded ; whitish- 
grey; cilia grey, apices paler. 

New South Wales: Mount Kosciusko (5,000 ft.), 
December ; three specimens. Type in Coll. Goldfinch. 


EUPHYIA TRISSOCYMA, N. sp. 
TpLoToKv/L0S, three times waved. 


gd, 22 mm. Head grey-whitish. Palpi 21; fuscous, 
whitish beneath. Antennae grey-whitish ; iltabiontt in male 4. 
Thorax grey-whitish ; patagia with a postmedian, transverse, 
fuscous line. Abdomen whitish with some fuscous irroration, 
and paired fuscous dots on some segments. Anterior legs 
fuscous [middle and posterior pairs missing]. Forewings tri- 
angular, costa nearly straight, apex round-pointed, termen 


= 

4 * 
Ay 

We 


953 


bowed, oblique, wavy; whitish with oblique, transverse, 
fuscous lines; a moderate fuscous basal patch, posterior edge 
from + costa to near base of dorsum; two very fine incomplete 
lines follow this; a broad, gently outwardly curved line from 
mid-costa to 4 dorsum; a dark-fuscous median discal dot; two 
very fine incomplete lines in median area; a broad threefold 
line from 3? costa to # dorsum, slightly bent outwards beneath 
costa, and again in middle; four very fine incomplete lines 
follow this; a well-marked terminal line, interrupted on veins ; 
cilia whitish, apices partly fuscous. Hindwings with termen 
slightly rounded, wavy; whitish; many fuscous lines from 
dorsum, more or less obsolete towards costa; terminal line and 
cilia as forewings. 

New South Wales: Jervis Bay, in October; one speci- 
men. Type in Coll. Goldfinch. 


-KUPHYIA APREPTA, NL. sp. 

dmpertos, undistinguished. 

@, 36 mm. Head and thorax fuscous. Palpi 21; fuscous, 
beneath ochreous-whitish towards base. Antennae fuscous. 
Abdomen fuscous with fine ochreous-whitish irroration. Legs 
fuscous. Forewings broadly triangular, costa moderately 
arched, apex rounded-rectangular, termen slightly bowed, 
slightly oblique, slightly crenulate; pale fuscous, basal patch 
and median band fuscous; basal patch small, posterior edge 
transverse, outwardly curved, wavy; two or three obscure lines 
precede median band; median band with anterior edge from 
4 costa to 4 dorsum, slightly outwardly curved, finely dentate ; 
posterior edge from % costa, at first nearly transverse, crenu- 
late, below middle bent inwards, and again transverse to 
2 dorsum; in this band is a darker median discal dot, pre- 
ceded and followed by a wavy transverse line, best marked 
towards costa; several faint and obscure transverse lines 
beyond band ; a crenulate, whitish, subterminal line; a narrow 
fuscous terminal line; cilia pale fuscous with a darker median 
line. Hindwings with termen rounded, crenulate; pale grey 
without markings; cilia pale grey. 

Victoria: Kyneton, in December; one specimen. Type in 
Coll. Lyell. 

HKUPHYIA CONIOPHYLLA, 0. sp. 


koviopvAdos, with dusty wings. 


2, 30 mm. Head reddish-brown mixed with fuscous. 
Palpi 34; fuscous, base beneath whitish. Thorax pale grey, 


_anteriorly reddish tinged. Abdomen pale grey mixed with 


ochreous-whitish and fuscous, base of dorsum reddish tinged. 
Legs fuscous; tarsi obscurely annulated with whitish; anterior 


254 


coxae reddish tinged. Forewings triangular, costa gently 
arched, apex acute, termen slightly bowed, oblique; whitish 
irrorated with fuscous-brown, which forms indistinct lines; a 
subbasal line from 3 costa, at ‘first outwardly oblique, but bent 
soon after origin, " thence slightly curved to near base of 
dorsum ; antemedian line very indistinct; a fuscous discal dot 
beneath mid-costa; postmedian very slender, from 4 costa 
obliquely outwards, angled beneath costa and in middle, 
thence to # dorsum; a fairly broad fuscous-brown terminal 
band, its anterior edge suffused, containing a fine, whitish, 
wavy, submarginal line; cilia fuscous-brown with pale basal 
and postmedian lines. Hindwings with termen rounded, 
slightly wavy; whitish irrorated with fuscous-brown, more 
densely towards termen; a faint whitish submarginal line; 
cilia grey, bases and apices paler. Underside whitish with 
fuscous-brown irroration and discal dots on fore- and hind- 
wings. 

New South Wales: Mount Kosciusko (5,000 ft.), in 
March; one specimen. 


DIPLOCTENA PANTOEA, Turn. 


Queensland: National Park (3,000 ft.), in February and 
March; seven specimens (4 males and 3 females). These are, 
ut consider, conspecific with southern examples, though they 
agree ill with my description, the species being exceedingly 
variable. The structure of the male antennae is the same. 
National Park examples are distinctly green with well-defined 
basal patch and median band fuscous-brown, but the latter 
sometimes incomplete; minute white dots are sometimes pre- 
sent on the subterminal line, and one female has a white 
dorsal dot in median band. Some examples from Lorne and 
Ebor, though in poor condition, approach these closely, but 
most of the males from these localities have the forewings 
almost wholly fuscous-brown. 


XANTHORHOE SODALIATA. 
Q. Cidaria sodaliata, Wlk., Cat. Brit. Mus., xxv., 


p. 1410. 
3. Coremia divisata, Wlk., Cat. Brit. Mus., xxxv., 
p. 1682. , 


Q. Xanthorhoé subidaria, var. urbana, Meyr., Proc. 
Linn. Soc. N.S. Wales, 1890, p. 864. 


This bala was first given by Swinhoe (Cat. Oxf. 
Mus., il., p. 345), but he identified the species with Guenée’s 
cymaria. T believe that Guenée’s description clearly applies 
to one of the forms I still include under swbidaria, Gn. 
Whether these are really all conspecific is open to doubt, and 


255 


until the male genitalia have been examined and compared 
by a competent authority, this doubt is likely to continue. 

Sodalhata female is very distinct by its uniform dark 
suffusion; the male has a uniformly dark median band on 
forewing, without brown or purplish tinge, while the terminal 
area is paler or even whitish. From eastern examples of male 
substdaria I have little difficulty in distinguishing it, but some 
Western Australian examples (which may represent a third 
species) are very similar. | 

Northern Queensland: Atherton, Herberton, Townsville. 
Queensland: Eidsvold, Gayndah, Nambour, Brisbane, Strad- 
broke Island, Mount Tambourine, Killarney, Nanango, Stan- 
thorpe, Roma. New South Wales: Murwillumbah, Lismore, 
Glen Innes, Ebor, Sydney, Moruya. Tasmania: Hobart. 
Also from Norfolk Island. 


XANTHORHOE EPIA, N. Sp. 
nos, soft. 


3, 9, 29-34 mm. Head brownish-grey, sometimes partly 
reddish tinged. Palpi 3; brownish-grey. Antennae grey; 
pectinations in male 6. Thorax and abdomen grey. Legs grey ; 
posterior pair paler. Forewings triangular, costa nearly 
straight to %, thence arched, apex pointed, termen bowed, 
oblique; grey with numerous fine, oblique, fuscous, transverse 
lines, more or less reddish tinged; sometimes the lines and 
disc are wholly reddish; a small slightly darker basal patch; 
median band darker, moderately broad on costa and in middle, 
then narrowed to dorsum to half this breadth, anterior edge 
from 4 costa to beyond 4 dorsum, slightly curved, posterior 
edge from 2 costa to before # dorsum, very obtusely angled 
outwards in middle, sometimes a fuscous discal dot beneath 
costa before middle; cilia pale fuscous, reddish tinged, apices 
paler. Hindwings with termen rounded; grey; a series of 
alternate darker and paler transverse lines from dorsum not 
reaching middle; a fine, interrupted, fuscous terminal line; 
cilia grey. 

The sexes are similar. Nearest X. centroneura, Meyr., 
which has the ground-colour much paler and contrasting with 
the median band, whose outer edge is more angled, and has 
also numerous blackish dots on veins. 

New South Wales: Mount Kosciusko (5,000 ft.), in 
February and March; 5 male and 6 female examples. 


XANTHORHOE METOPORINA, 0. sp. 
perorwpivos, autumnal. 


‘ Q, 32 mm. Head grey-whitish with dark fuscous; tuft 
fuscous. Palpi 24; fuscous; base narrowly white. Antennae 


256 


grey. Thorax and abdomen grey. Legs fuscous, irrorated, 
and tarsi annulated with whitish. Forewings broadly tri- 
angular, costa moderately arched, apex round-pointed, termen 
straight, oblique, crenulate; brown-whitish; markings fus- 
cous; a moderate basal patch, its posterior edge well defined, 
obliquely rounded, from % costa to § dorsum; a moderately 
broad median band, anterior edge outwardly curved, ill- 
defined, from 4 costa to 4 dorsum, posterior edge from 3 costa 
to 3 dorsum, with a large acutely-angled median projection ; 
several very fine, ill-defined, finely-waved lines precede and 
follow median band, and are traceable in the band itself; a 
dark-fuscous discal dot slightly before middle; a fine terminal 
line; cilia fuscous, bases and apices partly whitish. Hind- 
wings with termen gently rounded, crenulate; pale grey, with 
indications of fine, transverse, fuscous lines towards dorsum ; 
cilia grey, bases and apices partly whitish. Underside fuscous- 
grey, with dark-fuscous discal dots on fore- and hindwings. 

New South Wales: Mount Kosciusko, on March 2, 1912; 
two specimens. ; 

Gen. DASYSTERNICA, n. gen. 

I substitute this name for Dasysterna, Turn., which is 

preoccupied. 


DASYSTERNICA PERICALLES, Nl. Sp. 


mepikadAys, very beautiful. 


3,9, 23-27 mm. Head dark fuscous irrorated with 
ochreous. Palpi 3; ochreous with some dark-fuscous hairs. 
Antennae dark fuscous with fine whitish annulations; in male 
thickened and slightly laminate, ciliations +. Thorax dark 
fuscous irrorated with ochreous. Abdomen dark fuscous plen- 
tifully irrorated with ochreous; beneath ochreous. Legs pale 
ochreous with fuscous irroration, tarsi fuscous annulated with 
pale ochreous. Forewings triangular, costa slightly arched, 
apex round-pointed, termen bowed, oblique; fuscous with 
brownish and whitish irroration in parts; a basal patch limited 
by an outwardly curved, dark-fuscous and brownish, trans- 
verse, subbasal fascia; beyond this is a pale fascia containing 
some whitish irroration; median band outlined by two dark- 
fuscous and brown fasciae, its centre paler, with a minute, 
fuscous, median, discal dot sometimes indicated; anterior 
fascia from 4 costa to 2 dorsum, outwardly curved, its anterior 
edge twice indented and whitish ; posterior fascia from % costa 
to # dorsum, its posterior edge whitish, with a small posterior 
tooth above middle, and a large bidentate prominence in 
middle; one or two fine parallel fuscous lines beyond this are 
sometimes traceable; sometimes an indistinct pale subterminal] 
line; cilia fuscous, apices whitish-ochreous or barred with 


a 


257 


whitish-ochreous. Hindwings with termen rounded; orange; 
some fuscous irroration at base; three fine, fuscous, transverse 
lines, strongly angled in middle, in female obsolete; a dark- 
fuscous terminal band, much narrower in female; cilia as 
forewings. Underside ochreous; forewings with fuscous discal 
dot, postmedian fascia strongly dilated towards dorsum so as 
to join a terminal fascia, which is, however, mostly obsolete 
in female; hindwing with postmedian line and terminal fascia. 
in male, in female hardly developed. 

Tasmania: Cradle Mountain, in January; two specimens 
received from Dr. R. J. Tillyard. Type in Coll. Lyell. 


DASYSTERNICA CRYPSIPHOENA, 0. sp. 
Kpuiudowos, with hidden red. 


@, 26 mm. Head and palpi dark fuscous irrorated with 
whitish. Antennae fuscous. Thorax dark fuscous irrorated 
with whitish. [Abdomen broken off.] Legs dark fuscous 
irrorated, and tarsi annulated, with whitish [posterior pair 
missing]. Forewings triangular, costa slightly arched, apex 
round-pointed, termen bowed, slightly oblique, slightly crenu- 
late; whitish suffused with grey and on costa with fuscous; 
a subbasal fuscous fascia, containing some reddish scales, not 
reaching dorsum; this is followed by a whitish line, and this 
again by a ferruginous fascia at %, becoming fuscous at ex- 
tremities, and containing a small patch of reddish scales 
beneath costa; a median band consisting of two fasciae 
enclosing a pale area in which is a minute, fuscous, median, 
discal dot; inner fascia at 4, outwardly curved, edged with 
fuscous and partly filled in with reddish-ferruginous; outer 
fascia from # costa at first outwardly oblique, with an 
obtusely-angled posterior projection beneath costa, and 
another, double, in middle, thence dentate to 2 dorsum, out- 
lined with dark fuscous, and containing some reddish streaks 
on veins; a reddish-ferruginous band of suffusion separated 
from preceding fascia by a whitish line, and containing a 
wavy fuscous line; some obscure fuscous spots on termen; 
cilia fuscous barred with whitish. Hindwings with termen 
rounded ; grey, with three obscure whitish lines beyond middle, 
parallel to termen; cilia grey, apices whitish. Underside of 
forewings paler than upperside, with four transverse fuscous 
lines, the first median, the second followed by a whitish line, 
the fourth by a series of whitish dots; of hindwings like that 
of forewings, with a discal fuscous dot at 4. 

Type in Queensland Museum. It is possible that this 
may be identical with Lyirrhoé bertha, Swin. (Trans. Ent. 
Soc., 1902, p. 648). 


258 


Tasmania: Mount Wellington, in January; one specimen 
received from Mr. G. H. Hardy. 


DasyvrRis MELANCHLAENA, Nl. Sp. 
peAayxAavos, black-cloaked. 


3, Q, 24-28 mm. Head and thorax blackish, sometimes 
with a few whitish scales. Palpi 4; covered with long dense 
blackish hairs. Antennae blackish; ciliations in male im- 
perceptible. Abdomen blackish; some whitish scales on apices 
of segments. Legs blackish. Forewings triangular, costa 
slightly doubly sinuate, apex round-pointed, termen bowed, 
oblique; dark fuscous with obscure indications of darker 
transverse lines; and a few scattered whitish scales; an in- 
complete, very slender, outwardly curved, whitish, transverse 
line at 4; a better-marked whitish line from 4 costa to 2 
dorsum, slightly curved outwards and dentate; a dark-fuscous 
median discal dot outlined with whitish, postmedian whitish, 
from ? costa to # dorsum, sinuate, subdentate; an inter- 
rupted whitish subterminal line; cilia dark fuscous. Hind- 
wings with termen rounded; dark fuscous; sometimes a 
terminal series of whitish dots on veins, cilia dark fuscous. 

New South Wales: Mount Kosciusko (5,000 ft.), in 
December ; four specimens. Type in Coll. Goldfinch. 


ADDITIONAL LOCALITIES. 


Sauris hirudinata, Gn.—N. Q’land: Herberton; Q’land: Nam- 
bour, Blackbutt, Mount Tambourine, National Park (2-3,000 
ft.), Toowoomba; N.S. Wales: Lismore, Gosford. 

S. lichenias, Meyr.—N. Q’land: Herberton; Q’land: Toowoomba. 

Euchoeca rubropunctaria, Dbld.—Q’land: Coolangatta; N.S. 
Wales: Ebor, Nowra. 

Poecilasthena thalassias, Meyr.—N. Q’land: Herberton; Q’land: 
Gayndah, Coolangatta, National Park (2-3,000 ft.), Too- 
woomba, Bunya Mountains, Stanthorpe. 

P. pulchraria, Dbld.—N. Q’land: Herberton; Q’land: Stradbroke 
Island, Bunya Mountains (3,500 ft.), Stanthorpe; N.S. Wales: 
Lismore, Ebor; Vict.: Beaconsfield; Tas.: Tasman Peninsula; 
W. Austr.: Bridgetown, Perth. ‘ 

P. balioloma, Turn.—N.S. Wales: Glen Innes; Vict.: Mount St. 
Bernard (5,000 ft.). 

P. glaucosa, Luc.—Q’land: National Park (2-2,500 ft.). 

Minoa euthecta, Turn.—Q’land: Gayndah, Toowoomba, Bunya 
Mountains (3,500 ft.), Killarney. 

Gymnoscelis delocyma, Turn.—N. Terr.: Darwin. 

G. acidna, Turn.—N. Q’land: Cooktown, Cairns. 

G. mesophaena, Turn.—N. Q’land: Herberton. 

G. callichlora, Turn.—N. Q’land: Herberton. 

G. aenictopa, Turn.—N. Q’land: Herberton. ; 

Chloroclystis catastreptes, Meyr.—Q’land: Nambour, National 
Park (3,000 ft.), Toowoomba, Bunya Mountains (8,500 ft.); 
N.S. Wales: Katoomba, Nowra. 


259 


C. testulata, Gn.—Q’land: Toowoomba; N.S. Wales: Ebor, 
Mount Kosciusko ; Vict.: Castlemaine. 

. insigillata, Wlk.—Q’land: Toowoomba; N.S. Wales: Ebor, 
Mount Kosciusko. 

C. approximata, Wlk.—N. Q’land: Cairns, Herberton; Q’land: 
Mount Tambourine, National Park (8, 000 ft. ye Ne S. Wales: 
Lismore. 

C. laticostata, Wlk.—Q’land: Gayndah, Mount Tambourine, 
Coolangatta, National Park (8, 000 ft. Vi, Toowoomba, Killarney, 
Roma, Charleville ; N.S. Wales: Lismore, Ebor, Nowra, 
Adaminaby : Vict.: Beaconsfield, Daytrap; W. Austr.: 
Busselton, Perth. 


2 


arene aa aaa 


t pyrrholopha, Turn.—N. Q’land: Atherton, Herberton. 
metallospora, Turn.—Q’land: Gayndah. 

. cissocosma, Turn.—N. Q’land: Cairns, Herberton; Q’land: 
Nambour, National Park (3,000 ft.), Toowoomba. 
mniochroa, Porn =N. Q’land: Cairns, Atherton. 
gonias, Turn.—N. Q’land : Herberton ; Q’land: Stradbroke 
Island; N.S. Wales: Manning River. 

. alpnista, Turn.—N. Q’land: Herberton. 

. bryodes, Turn.—N. Q’land: Herberton; Q’land: Rosewood. 

. elaecopa, Turn.—N. Q’land: Herberton. 
athaumasta, Turn.—N. Q’land: Herberton. 
filata, Gn.—Vict.: Beaconsfield, Castlemaine; Tas.: Mount 
Wellington. 

C. leptomita, Turn.—Q’land: Brisbane, National Park (3,000 ft.). 


Tephroclystia melanolopha, Swin.—N. Q’land : Cairns, Herberton; 
Q’land: Nambour, Brisbane. 

Mnesiloba eupitheciata, Wik.—N. Q’land: Cairns, Herberton; 
_ Q’land: Nambour, Mount Tambourine, Southport, Too- 
woomba. 

Microdes villosata, Gn.—N.S. Wales: Nowra, Mount Kosciusko. 

M. squamulata, Gn.—N.S. Wales: Glen Innes; Vict.: Birchip; 
Tas.: Hobart. ; 

Chaetolopha oxyntis, Meyr.—N. Q’land: Cairns; Q’land: Mount 
- Tambourine, National Park (2-3,000 ft.); N.S. Wales: Lis- 
more, Sydney. 

Cc, leucophragma, Meyr.—Q’land: Nambour; N.S. Wales: Ebor; 
Vict. : Dunkeld. 

C. emporias, Turn.—N. Q’land: Herberton. 

C. niphosticha, Turn.—Q’land: Nationa] Park (3-4,000 ft.). 
Scotocyma albinotata, Wlk.—N. Q’land: MHerberton; Q’land: 
Nambour. . 
Eccymatoge callizona, Low.—N. Q’land: MHerberton; Q’land: 

Nambour, Brisbane; N.S. Wales: Glen Innes. 

Horisme peplodes, Turn.—Q’land: Caloundra, Toowoomba, Roma. 

H. scotodes, Turn.—N. Q’land: Herberton; Q’land: Caloundra; 
N.S. Wales: Port Macquarie, Nowra. 

EKucymatoge ghosha, Wlk.—N. Q’land: Herberton: Q’land: 
Caloundra, Stradbroke Island, Nationa] Park (3,000 ft.). 

EK. aorista, Turn.—N. Q’land: Innisfail, Herberton; Q’land: 
Blackbutt, Mount Tambourine; N.S. Wales: Lismore, 
Sydney. 

Heterochasta conglobata, Wlk.—N. Q’land: Cairns, Herberton; 
Q’land: Mount Tambourine, National Park (3,000 ft.); 
N.S. Wales: Dorrigo, Bulli. 


260 


Polyclysta hypogrammata, Gn.—N. Q’land: Atherton, Herberton ; 
Q’land: Stradbroke Island, National Park (3, 000 ft. ), Too- 
woomba, Bunya Mountains 13) 500 ft.); N.S. Wales: Lismore. 

Cidaria scythr opa, Meyr.—Q’land : Nambour, Caloundra, Too- 
woomba, Bunya Mountains; N.S. Wales: Lismore. 

C. lasioplaca, Low.—N. Q’land: Herberton; Q’land: Nambour, 
Toowoomba; N.S. Wales: Lismore. , 

. mMicrocyma, Meyr. —Tas.: Tasman Peninsula. 

. uncinata, Gn.—S. Austr.: Adelaide. 

. subochraria, Dbld. —Q’land: Killarney, National Park (3,000 
ft.); N.S. Wales: Ebor, Mount Canoblas, Moruya, Mount 
Kosciusko, Adaminaby ; Vict. : Moe, Dunkeld. 

Larentia epicrossa, Meyr.—Tas. : Cradle Mountain. 

L. dascia, Turn.—N.S. Wales: Sydney ; Tas.: Tasman Peninsula. 

Melitulias glandulata, Gn.—N.S. Wales: Mount Kosciusko (5,000 
ft.); Tas.: Mount Wellington. 

Euphyia phaedra, Meyr.—Q’land: Caloundra, Killarney; N.S. 
Wales: Murwillumbah. 

E. interruptata, Gn.—N.S. Wales: Mount Kosciusko (3-3,500 ft.). 

_ epicteta, Turn.—Tas.: Cradle Mountain. 

i ihnenee: Meyr. —Vict.: Castlemaine. 

‘ lucidulata, Wl1k.—N.S. Wales: Ebor; Vict.: Moe; Tas.: 

Tasman Peninsula. 

. conifasciata, Butl.—N.S. Wales: Ebor, Mossvale, Mount 

Kosciusko (5,000 ft.). 

percrassata, Wlk.—N.S. Wales: Mount Kosciusko (5,000 ft.). 

subrectaria, Gn.—Q’land: Mount Tambourine, Rosewood, 

Stanthorpe; N.S. Wales: Glen Innes, Ebor; Vict.: Moe. 

anthracinata, Gn.—Vict.: Melbourne; Tas.: Cradle Moun- 

tain, Mount Wellington. 

strumosata, Gn.—N.S. Wales: Ebor, Sydney; Tas.: Mount 

Wellington. 

vacuaria, Gn.—N.S. Wales: Mount Kosciusko (3,500-5.000 ft.) ; 

Vict.: Mount St. Bernard (5,000 ft.); Tas.: Cradle Mountain. 

symphona, Meyr.—Vict.: Mount Erica. 

excentrata, Gn.—Q’land: Killarney; N.S. Wales: Lismore, 

Armidale, Ebor. 

aglaodes, Meyr.—Vict.: Mount St. Bernard (5,000 ft.). 

. Imperviata, Wlk.—Vic.: Timberoo; S. Austr.: Adelaide ; 

W.A.: Perth. 

. heteroleuca, Meyr.—Vict.: Mount St. Bernard. 

languescens, Rosen.—N.S. Wales: Mount Kosciusko (5,000 ft.). 

. polycarpa, Meyr.—Tas.: Cradle Mountain. 

. chrysocyma, Meyr.—Tas.: Cradle Mountain. 

. perornata, Wlk.—Tas. : Cradle Mountain. 

insulsata, Gn.—Vict.: Dunkeld. 

mecynata, Gn.—Q’ land: Toowoomba; N.S. Wales: Glen Innes, 

Ebor, Taree, Mount Kosciusko (3- 3, 500 ft.); Vic.: Dunkeld. 

; polyxantha, Meyr.—N.S. Wales: Ebor; Vict.: Mount 

Macedon. 

. trygodes, Meyr.—N.S. Wales: Ebor. 

. severata, Gn.—Q’land: Toowoomba; N.S. Wales: Nowra; 

W. Austr.: Perth. 

. squamulata, Warr.—Vict.: Castlemaine. 

. opipara, Turn.—N.S. Wales: Mount Kosciusko (5,000 ft.). 

. ptochopis, Turn.—N.S. Wales: Moruya. 

iploctena argocyma, Turn.—N.S. Wales: Mount Kosciusko 
(5,000 ft.); Vict.: Mount St. Bernard. 


Qo@ 


gees Bi Javidi: Be Se Se eS Se & eee 


261 


Xanthorhoé subidaria, Gn.—Q’land: Clermont. 

X. brujata, Gn.—N. Q’land: Atherton, Herberton; Q’land: 
Gayndah, Stradbroke Island, Mount Tambourine, Coolangatta, 
National Park (3,000 ft.); N.S. Wales: Lismore, Glen Innes, 
Ebor; Vict.: Moe. | 

X. anaspila, Meyr.—Q’land: Brisbane, Toowoomba, Stanthorpe; 
N.S. Wales: Ebor, Mount Kosciusko (5,000 ft.).; Tas.: Mount 
Wellington. 

X. heliacaria, Gn.—N.S. Wales: Mount Kosciusko. 

X. vicissata, Gn.—Vict.: Beaconsfield, Moe, Dunkeld. 

Dasyuris decisaria, Wlk.—Vict.: Castlemaine. 

D. euclidiata, Gn.—N.S. Wales: Glen Innes, Ebor, Adaminaby. 

D. hedylepta, Turn.—N.S. Wales: Mount Kosciusko (5-6,000 ft.). 


Fam. ACIDALIADAE. 
EoIs FERRILINEA, Warr. 
E. cletima, Turn. 

Having now a good series of this species I find that the 
character on which I relied for the distinction of EF. cletima, 
the absence of an acute subcostal projectiom on postmedian 
line of forewing, is not trustworthy; this line varies in form. 


Northern Territory: Darwin. North Queensland : Towns- 
ville. Queensland: Duaringa, Gayndah, Brisbane, Stan- 


_thorpe. New South Wales: Sydney. 


Eo1s costaria, Wlk. (Acidalia). 
Acidalhia albicostata, Meyr. 


Queensland: Duaringa, Brisbane, Stradbroke Island, 
Coolangatta, Toowoomba, Stanthorpe, Chinchila, Charleville. 
New South Wales: Glen Innes, Sydney, Bathurst, Mount 
Kosciusko. Tasmania: Launceston, Deloraine. 


Eo1s ALBIcostaTa, Walk. 
Acidalia tsomorpha, Meyr. 
Eois costaria, Turn. 


While giving the wrong name to this species, I correctly 
pointed out the distinctions between it and the preceding. 
Not only are the posterior legs of the male quite different, 
but it is usually larger, more deeply pink, and the fillet is 
fuscous, not whitish or grey. 

Northern Territory: Darwin. Northern Queensland: 
Herberton. Queensland: Nambour, Brisbane, Stradbroke 
Island, Toowoomba, Stanthorpe. New South Wales: Tabu- 
lam, Glen Innes, Sydney. Victoria: Gisborne. ‘Tasmania: 
Hobart. South Australia: Mount Lofty. Western Aus- 
tralia: Waroona. 


262 


EKOIS MILTOPHRICA, 0. sp. 
pudtoppikos, rippled with red. 


Q, 18-20 mm. Head grey; face dark fuscous. Palpi 
scarcely 1; grey with a few dark-fuscous scales. Antennae 
pale grey. Thorax grey, with a minute, reddish, posterior 
dot. Abdomen grey with a median reddish dot on the dorsum 
of each segment except the first. Legs whitish; anterior pair 
grey. Forewings triangular, rather narrow, costa gently 
arched, apex round-pointed, termen bowed, oblique; grey with 
purple reflections; six rather broad, undulating, reddish- 
orange, transverse lines; first subbasal, incomplete, indicated 
only towards dorsum; second from 4 costa to 4 dorsum; third 
from mid-costa to beyond mid-dorsum; fourth from 4. costa 
to tornus; fifth from 2 costa to termen above tornus; sixth 
near termen meeting fifth; cilia grey. Hindwings with termen 
rounded ; as forewings but with only five red lines. Under- 
side grey with three darker postmedian lines on each wing. 

Although the male is unknown, this species may be 
easily recognized by its red lines. 

Northern Territory : Darwin, in November and December ; 
four specimens received from Mr. F. P. Dodd. 


KOIS SCAURA, 0. sp. 


scaurus, club-footed. 


36, 9,18mm. Head pale grey; collar and face fuscous. 
Palpi about 1; pale giey, upper-surface towards apex fuscous. 
Antennae grey; ciliations in male 1}. Thorax and abdomen 
pale grey. Legs pale grey; posterior pair ochreous-whitish ; 
posterior tibiae of male thickened, longer than femora, with 
a large expansile tuft of long hairs from base, without spurs, 
tarsi thickened, aborted, about +; of female normal but with: 
terminal spurs only. Forewings triangular, rather narrow, 
costa straight to middle, thence arched, apex round-pointed, 
termen straight, oblique; pale grey; faintly darker, dentate, 
transverse lines, which are minutely dotted with dark fuscous 
and pale edged posteriorly, at +, middle, and 4; a fine, wavy, 
pale, subterminal line; an interrupted dark-fuscous terminal 
line or series of dots; cilia pale grey. Hindwings with termen 
rounded ; as forewings. 

Near F. eretmopus but greyer, the male posterior tibiae 
are similar, but the tarsi much smaller and not dilated into 
paddle-shaped organs. 

Northern Queensland: Herberton, in November and 
January; three specimens (1 male and 2 females) received 
from Mr. F. P. Dodd. 


263 


| Eois EPicyrta, Turn. 
New South Wales: Mount Kosciusko (3,500 ft.). 


EoIs ELACHISTA, 0. sp. 
éAaxiotos, very small. 


3, 9, 12-13 mm. Head ochreous-whitish; face dark 
fuscous. Palpi under 1; fuscous. Antennae ochreous-whitish ; 
in male with tufts of long ciliations (3); in female slightly 
serrate. Thorax and abdomen ochreous-whitish. Legs ochreous- 
whitish; posterior pair in male very short, tibiae longer 


_ than femora, slightly thickened with scales on upper-surface, 


without spurs, tarsi 4; in female with terminal spurs only. 
Forewings rather broadly triangular, costa straight to %, 
thence arched, apex rounded, termen scarcely bowed, oblique ; 
ochreous-whitish with. a few dark-fuscous scales; a dark- 
fuscous dot on 4 costa; first line obsolete; a blackish discal 
dot beyond middle; a second dark-fuscous dot on 4 costa, 
from which proceeds a very slender, nearly obsolete, out- 


' wardly curved line, angled inwards above dorsum, ending on 


3 dorsum; some minute terminal dark fuscous dots; cilia 
ochreous-whitish. Hindwings with termen strongly rounded ; 
ochreous-whitish with a few dark-fuscous scales; lines obsolete ; 
a blackish discal spot before middle; cilia ochreous-whitish 
with a series of minute, subbasal, dark-fuscous dots. 

Nearest #. elaphrodes. The antennal structure of male 
furnishes a good character. 

Northern Territory: Darwin, in November; three speci- 
mens (1 male and 2 females) received from Mr. F. P. Dodd. 


Kois cHioristis, Meyr. (Acidalia). 

This must be an Hows. Meyrick states that 6 and 7 of 
hindwings are stalked. I have a female from Caloundra, 
Queensland, with terminal spurs only on posterior tibiae, 
to which I refer here, but unfortunately no male. The follow- 
ing species is closely allied. | 


EOIS PRIONOSTICHA, 0. sp. 


mpiovootixos, With saw-like line. 


3, 9, 19-22 mm. Head white; collar and face fuscous. 
Palpi under 1; fuscous or fuscous-whitish. Antennae grey ; 
in male with tufts of moderately long cilia (14). Thorax and 


abdomen white. Legs whitish; anterior pair fuscous in front; 


posterior pair in male short, tibiae much longer than femora 
(14), smooth, dilated towards apex, without spurs, tarsi very 
short ($); in female with terminal spurs only. Forewings 
triangular, costa straight to near apex, apex round-pointed, 


264 


termen slightly bowed, slightly oblique; white without 
ochreous tinge; a few scattered blackish-scales and a blackish 
discal dot beyond middle; lines grey; first from } dorsum, | 
obsolete towards costa; second from mid-costa, irregularly 
dentate, curving inwards in a short incomplete circle round 
discal dot, ending on mid-dorsum; third from 3 costa, finely 
dentate, nearly straight, to # dorsum; fourth subterminal ; 
fifth slender, submarginal; an interrupted terminal line; 
cilia whitish. Hindwings with termen strongly rounded; as 
forewings but without first line; discal dot before middle, 
minute or absent. | 

Very similar to #. chloristis, but Meyrick states that the 
posterior tarsi of male in this species are 4; also to ZH. poly- 
gramma; but Lower states that in this the discal dot of 
forewings is just anterior to median line. 

Northern Territory: Darwin, in November; three speci- 
mens (1 male and 2 females) received from Mr. F. P. Dodd 
and Mr. G..F. Hill. 

EOIS ARGOPHYLLA, 0. Sp. 

dpyopuAXos, white-winged. 

@, 18-20 mm. Head with fillet grey, posteriorly edged 
by a transverse blaekish line; collar and face fuscous. Palpi 1; 
grey, anteriorly whitish. Antennae grey. Thorax and 
abdomen white. Legs whitish; anterior pair grey in front; 
posterior tibiae in female with terminal spurs only. Fore- 
wings triangular, costa gently arched, apex round-pointed, 
termen slightly. bowed, oblique; shining white; without discal 
dot or irroration; costal edge grey; three slender, finely 
dentate, grey, transverse lines; first from 4+ dorsum, obsolete 
towards costa; second from % costa, nearly straight, to dorsum 
beyond middle; third nearly straight, subterminal; an inter- 
rupted grey terminal line; cilia white. Hindwings with 
termen rounded; as forewings. 

Readily distinguished from the two preceding species by 
the colour of the head. 

Northern Queensland: Evelyn Scrub. near Herberton, in 
January; two specimens received from Mr. F. P. Dodd, of 
which one is in Coll. Lyell. 


EoIs DELOSTICTA, N. sp. 
dyAootixtos, plainly spotted. 


, 18 mm. Head ochreous-whitish ; face dark fuscous. 
Palpi slightly over 1; fuscous. Antennae ochreous-whitish. 
Thorax ochreous-whitish with a posterior dark-fuscous dot. 
Abdomen ochreous-whitish; first segment with two dark- 
fuscous dots, each remaining segment with one median dorsal 


265 


dot. Legs ochreous-whitish; anterior pair fuscous in front, 
posterior tibiae in female with terminal spurs only. Forewings 


triangular, costa gently arched, apex rounded, termen bowed, 


oblique; ochreous-whitish with slight pale-grey suffusion and 
dark-fuscous dots; a median basal dot; five dots representing 
an antemedian line angled outwards beneath costa; a median, 
subcostal discal dot; a series of dots in a line from 2 costa to 
mid-dorsum; another series representing an undulating sub- 
terminal line; some grey submarginal suffusion; a terminal 
series of dots extending into cilia; cilia ochreous-whitish. 
Hindwings with termen rounded; as forewings. Underside 
similar. 


Northern Queensland: Kuranda, in June; one specimen. 


Gen, AcripauiA, Treit. 


I adopt this name for the genus to which I formerly 
attributed the name Leptomeris, Hb. The absence of long- 
stalking of veins 6 and 7 of the hindwings niay yenerally be 
relied on as a distinguishing character from Lois, though 
short-stalking is not uncommon. 


ACIDALIA DESPOLIATA, WIk. 


g6,18mm. Antennae moderately ciliated (1). Posterior 
femora of male short, tibiae elongate (24), swollen, smooth-. 
scaled, without spurs, tarsi very short in comparison (1/10th). 
No doubt the tibiae contain an internal groove and tuft of 
hairs which are not visible in my example. The relative sizes 
of femora, tibiae, and tarsi here attain their maximum dlis- 
proportion. A. optivata, which comes next, has tibiae 2, 
tarsi +. 

I took one male at Caloundra, Queensland, in October. 
. Northern Queensland: Cairns; one female in Coll. Lyell. 
Queensland: Stradbroke Island. 


ACIDALIA HYPOCHRA, Meyr. 
Acidalia axiotis, Meyr. 

I have received specimens from Western Australia, which 
differ in no way from those from Queensland. 

Northern Territory: Darwin. Northern Queensland: 
Thursday Island, Cooktown, Cairns, Herberton, Townsville, 
Ravenswood. Queensland: Duaringa, Gayndah, Nambour, 
Brisbane, Stradbroke Island, Southport, Coolangatta, Rose- 
wood. New South Wales: Sydney, Moruya. South Aus- 
tralia: Mount Lofty. Western Australia: Perth, Mundaring, 
York, Geraldton. Also from Norfolk Island. 


266 


ACIDALIA TENUIPES, Turn. 
Northern Territory: Melville Island. 


ACIDALIA SYNETHES, HN. sp. 
.ouvnOys, akin. 


g, 30 mm. Head pale grey; fillet white; face blackish. 
Palpi about 1; grey-whitish becoming dark fuscous towards 
apex. Antennae grey-whitish; in male serrate, ciliations 24. 
‘Thorax and abdomen pale grey. Legs pale grey; posterior 
_ pair in male whitish, tibiae dilated, tarsi 4. Forewings tri- 
angular, costa gently arched, apex tolerably pointed, termen 
slightly bowed, slightly oblique; pale grey without irrora- 
tion ; a dark-fuscous, subcostal, median, discal dot; lines very 
faintly marked ; antemedian line obsolete or nearly so; a very 
slender, finely dentate, sinuous line from % costa to 2 dorsum, 
a similar line from # costa to # dorsum, forming 
minute dots on veins; a very faint, whitish, dentate, 
subterminal line; a terminal series of fuscous interneural 
dots; cilia pale grey. Hindwings with termen rounded; as 
forewings but some grey irroration towards base, discal dot 
at 4, lines even less distinct. ; 

Very like A. otis, Meyr., from Mount Kosciusko, but 
greyer in colour, without any fuscous irroration, and posterior 
tarsi of male rather shorter relatively to tibiae. Type in 
Coll. Lyell. 


Western Australia: Waroona, in January; one specimen 
received from Mr. G. F. Berthoud. 


ACIDALIA PERIALURGA, 0. sp. 
mepiadoupyos, dyed with purple all round. 


9, 29 mm. Head grey; fillet white; face dark fuscous. 
Palpi 14; whitish becoming fuscous towards apex. Antennae 
grey, towards base whitish. Thorax grey. Abdomen grey- 
whitish sparsely irrorated with fuscous. Legs grey; posterior 
pair and middle femora ochreous-whitish with slight fuscous 
irroration. Forewings triangular, costa gently arched, apex 
round-pointed, termen bowed, oblique; grey with a few scat- 
tered fuscous scales; some pale-purplish suffusion towards 
base; a minute, fuscous, median, discal dot beneath costa; 
a band of pale-purplish suffusion, its inner edge from ¢ costa 
to # dorsum, slightly curved inwards above dorsum, outer 
edge formed by a fine, crenulate, fuscous line at about {, 
thickened to form minute dots on veins; a terminal series 
of dark-fuscous interneural dots; cilia pale purple with a 
few fuscous scales, apices grey-whitish. Hindwings with 


267 


termen slightly angled on vein 4; as forewings but discal 
spot at 4 and larger. 

New South Wales: Port Macquarie, in March, one speci- 
men. Type in Coll. Lyell. 


STERRHA OOPTERA, 0. Sp. 

@otTepos, oval-winged. 

Q,23 mm. Head whitish; face grey. Palpi about 1; grey. 
Antennae whitish-grey. Thorax and abdomen whitish-grey 
with slight grey irroration. Legs ochreous-whitish irrorated 
with grey; posterior tibiae with terminal spurs only. Fore- 
wings elongate-oval, costa gently arched, apex pointed, termen 
bowed, strongly oblique; whitish-grey irrorated with dark 
grey; a small, circular, fuscous, discal spot at 2; a fine, 
interrupted, dark-grey line from costa just before apex to 
% dorsum ; a similar terminal line; cilia whitish with two lines 
of grey irroration. Hindwings suboval, narrow, termen very 
strongly rounded; as forewings but discal spot median, and 
posterior line strongly curved. 


A curious-looking species, more suggestive of the genus 
Pylarge than Sterrha. 


Queensland : haicae one specimen received from Dr. 
Hamilton Kenny. 


STERRHA EUCLASTA, Ni. &p. 
Bp wee fragile. 


6, 24-26 mm. Head brown; fillet broadly white; face 
fuscous-brown. Palpi about 1, curved upwards, thickened 
with rough scales, terminal joint short; whitish. Antennae 
white; in male with fine short pectinations (4), ending in tufts 
of. long cilia (3). Thorax and abdomen ochreous-whitish. 
Legs fuscous ; posterior pair ochreous-whitish ; posterior tibiae 
of male with terminal spurs only, otherwise normal. Fore- 
wings rather narrowly triangular, costa gently arched, apex 
pointed, termen bowed, oblique; ochreous-whitish with slight 
grey suffusion and a very few fuscous scales; a minute fuscous 
discal dot beneath mid-costa; a suffused, straight, grey line 
from 2 costa to mid-dorsum; a similar double subterminal line 
from apex; a third line close to terminal margin; a series 
of minute, interneural, fuscous, terminal dots; cilia ochreous- 
whitish. Hindwings with termen rounded ; ochreous- whitish ; 
a fuscous discal dot before middle; a straight grey line from 
apex to ? dorsum; a faint parallel line posterior to this; 
terminal dots and cilia as forewings. 

New South Wales: Mount Kosciusko (3,500 to 5.000 ft.), 
= January; three specimens, of which one 1s in Coll. Gold- 

nch. 


268 


PROTOTYPA DRYINA, Turn. 
New South Wales: Ebor Scrub (4,000 ft.). 


CHRYSOCRASPEDA CRUORARIA, Warr. (Chrysolene). 
Chrysocraspeda aurvmargo, Warr. 
Chrysocraspeda inundata, Warr. 


I formerly regarded these as distinct. Mr. F. P. Dodd 
first pointed out to me that they are forms of one very variable 
species. 

Northern Queensland: Cooktown, Cairns. Also from 
New Guinea. 


GNAMPTOLOMA CHLOROZONARIA, Walk. (Thalassodes). 


This name supersedes mundissima, W1k. 

Northern Queensland: Cairns. Queensland: Duaringa, 
Bundaberg, Hidsvold, Gayndah. Also from Ceylon, India, 
and Africa. 

PERIXERA FLAVIRUBRA, Warr. 


2, 36 mm. Head brown; face whitish-ochreous with a 
purple transverse bar near upper edge. Palpi 3, terminal 
joint 4; purple, lower edge whitish-ochreous. Antennae, 
upper-surface fuscous, lower-surface ochreous-whitish. Thorax 
brown. Abdomen brown; towards apex pale grey; under- 
surface whitish-ochreous. Legs whitish-ochreous. Forewings 
triangular, costa slightly arched, apex round-pointed, termen 
bowed, slightly oblique, slightly dentate; yellowish-brown 
finely strigulated with dark brown; three fuscous dots on 
veins representing a subbasal line; a median discal dot, white 
edged with dark brown; a bisinuate line of-fuscous dots 
from 2 costa to # dorsum ; sometimes a dark-fuscous blotch on 
this line above middle; a terminal series of fuscous dots; 
cilia brown. Huindwings with termen rounded, dentate; as 
forewings; discal dot at 4 (in one example crescentic) ; some- 
times a dark-fuscous tornal blotch. Underside pinkish-white, 
with a posterior line of fuscous dots. 

Northern Queensland: Cooktown, Cairns, Herberton. 


PERIXERA LAPIDATA, Warr. 


3, 9, 32-40 mm. Head whitish with a few dark-fuscous . 
scales on vertex; upper half of face brown. Palpi in male 
24, terminal joint 4; in female 24, terminal joint 1; fuscous 
or purple-fuscous, beneath whitish. Antennae whitish; in 
male with slight fuscous irroration, pectinations 8, apical 4 
simple. Thorax whitish with a few fuscous scales. Abdomen 
ochreous-whitish with a few fuscous or purple scales towards 
base of dorsum. Legs ochreous-whitish; dorsum of first two 


269 


pairs and tuft on male posterior femora purple tinged. Fore- 
wings triangular, costa moderately arched, apex pointed, 
termen slightly bowed, oblique; whitish beset with numerous 
fine grey strigulae; subbasal line represented by three fuscous 
dots; a small, grey, pale-centred, discal spot before middle; 
a bisinuate, subterminal] line of fuscous dots; a terminal series 
of blackish interneural dots; cilia whitish. Hindwings with 
termen gently rounded, slightly dentate; as forewings, but 
without subbasal dots; discal spot at 4, larger, ochreous, out- 
lined with fuscous. Underside whitish with fuscous discal 
marks and subterminal series of dots. 

Northern Queensland: Cairns, Herberton. Also from 
New Guinea. 


ANISODES PULVERULENTA, Swin. 
Maculifera, Swin., and cyclophora, Turn., are the female 


of this species. 
Northern Queensland: Cairns, Herberton, Townsville 


Also from Malay Peninsula and India. 


PISORACA SIMPLEX, Warr. 


The species I have described as decretarra, Wlk., had 
better stand for the present under Warren’s name, as it is 
doubtful whether it is really Walker’s species. 


ADDITIONAL LOCALITIES. 


Mnesterodes trypheropa, Meyr.—Also from New Guinea. 
Xenocentris rhopalopus, Turn.—N,. Q’land: Herberton. 
X. pilosata, Warr.—N. Terr.: Darwin, Melville Island; Q’land: 
Rosewood. 
X. epipasta, Turn.—N.S. Wales: Lismore. 
Eois coercita, Luc.—Q’land: Nambour. 
. liparota, Turn.—Q’land: Rosewood. 
. eretmopus, Turn.—Q’land: Gayndah, Coolangatta. 
. plumbiscriptaria, Christ.—Q’land: Eidsvold. 
. halmaea, Meyr.—N. Q’land: Claudie River; Q’land: National 
Park (3, 000 "E, ); N.S. Wales: Ebor. 
. fucosa, Warr.—N. Terr.: Darwin. 
: philocosma, Meyr. — Q’land: Gayndah, Caloundra, Mount 
Tambourine, Coolangatta; N.S. Wales: Glen Innes. 
Acidalia lydia, Butl. —Q’land: Caloundra, Jandowae, Charleville; 
Vict.: Brentwood, Birchip; S. Austr. : Wynbring. 


ES See 


A. perlata, WIk. ep ead: National Park (2- 3,000 ft. a Killarney ; 
N.S. Wales: Ebor, Bega, Mount Kosciusko (5,000 ft.). 

A. liotis, Meyr.—Vict. : Mount St. Bernard (5,000 ft.). 

A. desita, Wlk.—N. Terr.: McDonald Ranges; N. Q’land: Her- 
berton ; Q’ land: Blackbutt, Rosewood. 

A. rubraria, Dbld.—Q’land: Eidsvold, Gayndah, Rosewood, Cool- 


angatta, Roma, Charleville, Cunnamulla: N.S. Wales: "Bega; 
Vict. : Gisborne, Birchip; "W. Austr. : Perth, Bridgetown. 


4 
v 
‘ 


. sublinearia, Wlk.—N. Terr.: Darwin; Q’land: Coolangatta; 
N.S. Wales: Sydney. 
. prosoeca, Turn.—N. Terr.: Darwin; Q’land: Hidsvold. 
. recessata, Wlk.—N. Q’land: Herberton; Q’land: Eidsvold, 
Gayndah, Rosewood. 
. nictata, Gn.—N. Q’land: Cairns, Ingham. 
. oppilata, Wik.—Q’land : Eidsvold, Gayndah, Stanthorpe, Roma, 
Charleville; N.S. Wales: Tabulam. 
. thysanopus, Turn.—N. Terr.: Darwin; N. Q’land: Horbentee ; 
Q’land: Killarney. 
. optivata, Wlk.—N. Q’land: Cairns, Atherton, Herberton; 
Q’land: Eidsvold, Gayndah, Coolangatta, Warwick, Killarney, 
Roma; N.S. Wales: Tabulam, Armidale, Ebor Bega; Vict. : 
Birchip; W. Austr.: Harvey, Busselton, Perth. 
A. caesaria, Wlk.—N. Terr. : Darwin; Q’ land: Stradbroke Island. 
Dasybela achroa, Low.—Vic.: Sale. 
Somatina maculata, Warr. ig! land: Hidsvold. 
Problepsis clemens, Luc.—Q’land: Toowoomba. 
P. sancta, Meyr. —Q’land: Blackbutt, Toowoomba. 
P. cana, Hmps.—N.W. Austr. : Derby. 
Ptychophyle cyphosticha, Turn.—N, Terr.: Darwin. 
Gnamptoloma aventiaria, Gn.—N. Q’land: Atherton, Herberton; 
Q’land: Emerald, Fidsvold, Gayndah, Caloundra, Rosewood ; 
N.S. Wales: Lismore. 

Organopoda olivescens, Warr.—N. Q’land: Herberton; Q’land: 
National Park (3,000 ft.). 

Brachycola obrinaria, Gn.—N. Terr.: Darwin. 

B. porphyropis, Meyr.—N. Q’land: Herberton; Q’land: Blackbutt, 
National Park (3,000 ft.); N.S. Wales: Lismore. 

Anisodes leptopasta, Turn.—N. Q’land: Cooktown. 

Pisoraca nephelospila, Meyr.—N. Q’land: Cooktown. 

P. punctata, Warr.—N. Q’land: Herberton. 

P. cryptorhodata, Wlk.—Q’land: Gayndah; N.S. Wales: Sydney. 


Fam. GEOMETRIDAE. 
Gen. IprocHRoa, n. gen. 


270 


> > PP PP Pp 


x 


idvoxpoos, with peculiar colouring. 


Frons flat. Tongue absent. Palpi minute (less than 4); 
porrect, shortly rough-haired. Antennae bipectinate in both 
sexes, extreme apex simple. Thorax and abdomen without 
crests; thorax not or very slightly hairy beneath. Posterior 
tibiae with two pairs of fully developed spurs; not dilated 
in male. Forewings with 7, 8, 9, 10 stalked from before 
angle of cell, 11 from cell, connected by a bar or anastomosing 
with 12. Hindwings with strong basal costal expansion, 
frenulum and retinaculum absent; 2 from middle of cell, 3 
from well before angle widely remote from 4, 6 and 7 connate 
or short-stalked, 8 touching cell at a point near base, thence 
very gradually diverging. , 

Near Cenochlora, Warr., but has two paits of spurs on 
posterior tibiae. Type I. demissa. 


271 


IDIOCHROA DEMISSA, ND. sp. 

demissus, modest. 

dg, 21-22 mm. Head green; face and palpi pale fuscous. 
Antennae whitish; pectinations in male 10, apical 4 simple. 
Thorax green. Abdomen whitish with a broad, dull-reddish, 
median, dorsal streak; beneath pale fuscous. Legs whitish- 
ochreous; anterior pair pale fuscous. Forewings triangular, 
costa gently arched, apex acute, termen slightly bowed, 
oblique; 11 connected with 12 by a long bar; rather dark 
green ; costal edge pale ochreous as far as middle; a fuscous 
dot on end of cell at about 2; cilia green. Hindwings with 
termen rounded; dull reddish; dorsum narrowly green; a 
darker reddish dot on end of cell; cilia whitish, slightly 
reddish tinged. Underside more or less suffused with dull 
reddish. 

Q@, 22 mm. Antennal pectinations 8. Face green. 
Hindwings pale green. Underside green. Differs from male 
in total absence of reddish colouring. 

Queensland: Rosewood, in September; Toowoomba, in 
December (W. B. Barnard); six specimens. 


IDIOCHROA CELIDOTA, Nl. sp. 
- «nXdwdwtos, blotched. 


36, 22 mm.; 9, 29 mm. Head white, posterior edge 
green ; face dark reddish. Palpi very short (about 4); red- 
dish. Antennae ochreous-whitish; pectinations in male 12, 
in female 6, extreme apex simple. Thorax green. Abdomen 
whitish tinged with reddish; dorsum of first two segments 
green; sometimes a suffused, fuscous, median, dorsal streak 
containing several white dots; under-surface ochreous-whitish. 
Legs whitish-ochreous; anterior pair reddish. Forewings tri- 
angular, costa gently arched, apex round-pointed, termen 
nearly straight, slightly oblique; 11 anastomosing with 12; 
green (inclining to bluish-green); costal edge pale ochreous; 
a large tornal blotch outlined with purple fuscous, whitish 
containing a pale-reddish streak along anterior border, and 
a broader pale-reddish central partition, in which are some 
purple-fuscous scales; cilia’ grey. Hindwinys with termen 
rather irregularly rounded, tornus rather prominent; colour 
and cilia as forewings, but without markings. Underside 
whitish-green; forewings ochreous tinged with a pale-grey 
tornal blotch. 

Queensland: Gayndah, female type received from Dr 
Hamilton Kenny; Rosewood, a wasted male, in April. 


272 


CyMATOPLEX HALCYONE, Meyr. (Hucrostes). 


This name supersedes crenulata, Luc. 

Northern Territory: Darwin. Northern Queensland: 
Thursday Island, Cairns, Townsville. Queensland: Caloundra, 
Brisbane, Stradbroke Island, Southport. Also from New 
Guinea. 


Gen. Mrxocera, Warr. 

This name supersedes Gynandria, Turn. Experience has 
shown me that pectination of the female antennae cannot be 
relied on as a generic character. The genus comes near 
Cymatoplex, but 11 arises from end of cell, connate with 
7, 8, 9, 10, or is short-stalked with them. In the latter 
genus 11 is from well before end of cell. Type M. parvulata, 
Wilk., from India. There are also five African species. 


Gen. Euvcrostses, Hb. 


Tongue weakly developed. Palpi slender, moderately long, 
porrect; terminal joint in male very short, m female longer. 
Femora smooth. Posterior tibiae without.middle spurs. Fore- 
wings with 3 and 4 connate, 5 from above middle, 6 from 
upper angle, 11 from cell, anastomosing with or running into 
12. Hindwings with cell short (2), with 3 and 4 connate, 
6 and 7 connate, 12 anastomosing with cell at a point near 
base, thence rapidly diverging. Frenulum and retinaculum 
absent and hindwings with costal expansion at base in both 
SEXES. 

Near Cymatoplex, Turn., and Mirocera, Warr. Differs 
from the first by the shorter cell of hindwing and rather 
longer female palpi; from the second by the origin of 11 of 
forewings well before end of cell. Type H. indigenata, De 
Villers, from the Mediterranean area. 


EUCROSTES IOCENTRA Meyr. 
Iodis barnardae, Luc. 

Mr. Prout makes Hucrostes nanula, Warr., a synonym; 
but I think Warren’s type is so wasted as to be unrecog- 
nizable. 

Queensland: Duaringa, Brisbane, Charleville. 


Gen. Ivtors, Prout. 


This genus has been made for argocrana, Meyr., a species 
which I have not seen. 


EULOXIA GRATIOSATA, Gn. 


I shall not follow Prout in placing this in a genus by 
itself under the name Mizochroa, Warr. The species occurs 


273 


rather commonly on Mount Kosciusko at 5,000 ft., with the 
oblique white line on forewing feebly developed or absent. 


EULOXIA ARGOCNEMIS, Meyr. (Jodis). 


Mr. Prout, who has doubtless examined the type, places 
it in this genus. 


CHLOROCOMA SYMBLETA, N. sp. 

ovupBAntros, comparable. 

3g, 36 mm. Head and face green; fillet broadly white. 
Palpi whitish, on upper-surface crimson. Antennae white, 
apical half and pectinations pale crimson; pectinations in 
male 5, apical 4 simple. Thorax bluish-green. Abdomen 
bluish-green; tuft, sides posteriorly, and under-surface 
whitish. Legs pale crimson; posterior pair whitish; posterior 
tibiae in male dilated with internal groove and tuft. Fore- 
wings broadly triangular, costa gently arched, apex sub- 
rectangular, termen very slightly bowed, moderately oblique; 
3 and 4 approximated at origin, 6 connate, 11 anastomosing 
with 12; bluish-green; costal edge white except near base 
and in apical +, where it is crimson; a darker green discal 
dot on end of cell; a very fine dentate whitish postmedian 
line obscurely indicated ; cilia pale crimson. Hindwings with 
termen rounded; 3 and 4 stalked; as forewings but without 
costal streak and discal dot. Underside pale green. 

Not unlike C. asemanta, Meyr., but this is a smaller 
species with green cilia. . 

New South Wales: Adaminaby (3,500 ft.), in October; 
one specimen. : 


CHLOROCOMA RHODOTHRIX, 0. sp. 

podofpré. rosy-haired. 

¢, 26 mm. Head and face brown; fillet broadly white. 
Palpi pale brown. Antennae white; pectinations fuscous 
[broken off except first two joints]. Thorax brown; posterior 
end and apices of patagia green. Abdomen green; tuft 
whitish; under-surface whitish-ochreous. Legs whitish ; 
anterior and middle pairs crimson anteriorly; both spurs on 


' middle tibiae and external spurs on posterior tibiae crimson ; 


posterior pair in male not dilated and without internal groove 
and tuft. Forewings triangular, costa straight except near 
base and apex, apex pointed, termen very slightly bowed, 
oblique; 3 and 4 connate, 6 short-stalked with 7, 8, 9, 10, 
11 anastomosing with 12; deep green; a broad brown costal 
streak from base to apex, leaving costal edge white from 
3 to #, and thence crimson; veins mostly faintly marked 
with pale crimson; termen narrowly crimson; cilia deep 


274 


crimson. Hindwings with termen strongly rounded; 3 and 4 
short-stalked; as forewings but without costal markings; a 
crimson antemedian discal dot on end of cell. Underside 
similar. 

- Tasmania: Cradle Mountain, in January (3,000-3,500 
ft.); one specimen, received from Dr. R. J. Tillyard. 


CHLOROCOMA MELOCROSSA, Meyr. 


I now regard'C. periphracta, Turn., as a well-marked 
local race of C’. melocrossa. I have found it only on Strad- 
broke Island, but examples intermediate between it and the 
typical form occur at Coolangatta, in both instances attached 
to Banksia serratifolia. 


CHLOROCOMA NEPTUNUS, Butl. 
Chloéres cissina, Turn. 


In describing this as a Chloéres I overlooked the very 
slender male frenulum, and minute retinaculum near to base 
of wing. 

Queensland : a py Gayndah, Rosewood, Too- 
woomba, Killarney. 


CHLOROCOMA TACHYPORA, Turn. 


Near the preceding but distinguishable by the white 
costal streak of forewings, and the face being not green but 
greenish-ochreous.. 

Queensland: Stradbroke Island, Southport. 


Gen. PaMPHLEBIA, Warr. 


Differs from Chlorocoma, Turn., in the forewings having 
vein 11 stalked from 10, and in the terminal joint of palpi 
being elongate in female. Type P. rubrolimbraria. 


PAMPHLEBIA RUBROLIMBARIA, Gn. (Amaurinia). 
Thalassodes diserta, Wk. 
Thalassodes simpliciaria, W1k. 
Nemoria rufotinctaria, Snel. 
Chlorocoma perigrapta, Turn. 


Northern Queensland: Ingham. Also from New Guinea, 
Borneo, Ceylon, and India. I am indebted to Mr. L. B. 
Prout. for the identification. 


Gen. GrELAsmMa, Warr. 
Prasinocyma, Warr. 


Type G. thetydaria, Gu., from India. I am unable to 
separate these two genera. Those species to which Gelasma is 


275 


restricted by Prout form a natural group, which embraces 
centrophylla, Meyr., calaina, Turn., epimitra described below, 
and orthodesma, Low. In both calaina and orthodesma the 
terminal joint of palpi in female is fully 4, and the only 
structural distinction appears to be the angling of the termen - 
of the hindwing on vein 4, which is insufficient. The genus, 
as I conceive it, is large but not unmanageable, comprising 
some 120 species. 


GELASMA ISERES, Nn. sp. 
ionpys, equally fitted. 
é, 30 mm. Head and face green; fillet broadly white. 
Palpi short (about 1); whitish. Antennae white; pectina- 


_ tions in male 10, whitish-ochreous. Thorax green. Abdomen 


green; apex and underside whitish. Legs pale ochreous; 
coxae whitish. Forewings triangular, costa straight to 3, 
thence gently arched, apex subrectangular, termen «nearly 
straight, slightly oblique; green with numerous, fine, 
whitish, minute, transverse strigulae; a white costal streak 
from near base to near apex; cilia green. MHindwings with 
termen bowed, tornus prominent; as-forewings but without 
costal streak. Underside whitish-green. 

Very like P. albicostata, which differs in the longer palpi 
(14) and whitish cilia. 

Northern Territory: Darwin, one specimen received from 
mc G. FE. Hill. 


GELASMA LYCHNOPASTA, Turn. (Prasinocyma). 
New South Wales: :Ebor Scrub (4,000 ft.). 


GELASMA EPIMITRA, 0. sp. 

émiutpos, girdled. 

36, 24 mm.; 9, 28 mm. Heéad bDluish-green; fillet 
white; face green. Palpi in male 14, terminal joint 4; in 
female 34, terminal joint 2; green; under-surface white. 
Antennae white, towards apex ochreous tinged. Thorax 
bluish-green. Abdomen bluish-green; tuft and under- 
surface white. Legs whitish; anterior pair green on dorsum. 
Forewings triangular, costa moderately arched, apex round- 
pointed, termen bowed, oblique; 11 free; bluish-green densely 
irrorated, except on two transverse fasciae, with lustrous 
whitish scales; first fascia moderate, at 4, indistinct towards 
costa; second fascia at #, narrow on costa, soon broadening 
and outwardly curved, then nearly straight and again nar- 
rower to dorsum, its anterior edge rather suffused, posterior 
edge sharply defined, crenulate; costal edge grey from } to 
apex; a blackish median discal dot; a green terminal line; 


276 


cilia pale green. Hindwings with termen angled on vein 4, 
wavy; as forewings but without first fascia; discal dot at 3. 
Underside pale green. 3 

Northern Queensland: Evelyn Scrub, near Herberton, in 
- January; female type received from Mr. F. P. Dodd. New 
South Wales: Mount Gregson, in March; one male in Coll. 
Goldfinch. 

GELASMA ORTHODESMA, Low. 
Northern Queensland: Cairns. Also from New Guinea. 


GELASMA CENTROPHYLLA, Meyr. 


Northern Queensland: Herberton. Queensland: Bris- 
bane, Stradbroke Island, Toowoomba. New South Wales: 
Sydney. Victoria: Melbourne, Beaconsfield, Gisborne. Tas- 
mania: George Bay, Kelso, Georgetown. 


Gen. CHRysocHLoROMA, Warr. 


This, though nearly allied to Gelasma, may be separated 
by the strong male frenulum, and the presence of a weak 
frenulum in female. It contains only the one Australian 
species and four from New Guinea. 


Gen. Eucrna, n. gen. 

evknAos, calm, tranquil. 

Frons flat. Tongue very weakly developed. Palpi short 
(slightly over 1), porrect; second joint with long rough hairs 
beneath ; terminal joint in female about 4, slender, pointed. 
Antennae in female simple. Thorax and abdomen without 
crests; thorax slighty hairy beneath. Posterior tibiae with- 
out middle spurs. Forewings with 2 from #, 3 from before 
angle remote from 4, 5 from above middle, 6 from angle, 
7, 8, 9, 10 stalked, 10 arising before 7, 11 anastomosing with 
12. Hindwings with strong, basal, costal expansion, frenulum 
and retinaculum absent in female; cell about 4, lower dis- 
cocellular oblique, costal edge of cell not much shorter than 
dorsal; 2 from %, 3 and 4 remote at origin, 6 and 7 connate 
or just stalked, 8 approximated to cell near base, thence 
gradually diverging. 

Unfortunately the male, which will probably show addi- 
tional characters, is unknown, and the true position of 
the genus remains uncertain. 


EUCELA AMALOPA, Nl. Sp. 


apadw7ros. soft-lookin g. 


9, 36 mm. Head and face green. Palpi and antennae 
whitish. Thorax green. Abdomen whitish with green dorsal 


SS 


277 


and sublateral streaks. Legs whitish; coxae and anterior 
femora green. Forewings triangular, costa nearly straight 
but arched towards base and apex, apex pointed, termen 
nearly straight, moderately oblique; rather pale green ; costal 
edge white; an outwardly curved white line from 4 costa 
to 2 dorsum; a white line, broad except towards costa, 
nearly straight, from 2 costa to mid-dorsum; cilia whitish. 
Hindwings with termen rounded; pale green; cilia whitish. 
Underside pale green with postmedian white line, preceded 
by a darker shade of green, on both wings. - 

New South Wales: Mount Kosciusko (5,000 ft.), in 
January; one specimen. 


METALLOCHLORA NEOMELA, Meyr. (Jodis). 
Pisina, Warr., and albolineata, Pagent., are synonyms. 
Northern Territory: Darwin. North-western Australia: 
Broome. . Also from New Guinea, New Britain, and 
Tenimber Island. 


Gen. Eucyciopes, Warr. 


I am unable to agree with Mr. Prout in separating all 
the species except buprestaria to form his new genus 
Amisozyga, for buprestaria is closely allied to them, the slight 
structural differences being merely specific. Mono-specific 
genera should only be made for species isolated by consider- 
able structural peculiarity ; on the other hand, comparatively 
slight structural characters, if definite and constant, may be 
useful in separating two nearly related groups of species. 


EUCYCLODES DENTATA, Warr. 


I now regard this as merely a female aberration of LZ. 
prerordes, Wk. 


AGATHIA OCHROTYPA, Nn. sp. 
3 
w®xpotumos, pale-marked. 


@, 40-42 mm. Head and thorax bright green. Palpi 
2, terminal joint 4; whitish, terminal joint fuscous. 
Antennae whitish-brown with some fuscous irroration. 
Abdomen bright green, beneath whitish. Legs whitish- 
brown ; anterior pair partly suffused with fuscous. Forewings 
triangular, costa strongly arched, apex rectangular, termen 
bowed, wavy, oblique; bright green with sparse, pale-grey, 


_ transverse strigulae; markings pale grey mixed with pale 


ochreous-brown ; costal edge pale grey with darker strigulae ; 
an ill-defined, small, subbasal fascia; a fascia from 4 dorsum, 
not quite reaching 4 costa, bent outwards in middle, some- 
what constricted above and below middle; a second fascia 


278 


commencing in a blotch beneath # costa, constricted beneath 
this, and again above % dorsum; cilia grey. Hindwings with 
termen wavy, produced to an acute angle on vein 4; as fore- 
wings but with basal and antemedian fasciae; postmedian 
fascia expanded towards dorsum; a fuscous-brown marginal 
dot above terminal projection, and a larger marginal spot 
bisected by a whitish line beneath projection; cilia whitish, 
on projection fuscous, towards tornus with a fuscous basal 
line. Underside green-whitish with indications of postmedian 
fasciae. 

Northern Queensland: Evelyn Scrub, near Herberton, 
in December and February; two specimens received from Mr. 


F. P. Dodd. 


HELICOPAGE CINEREA, Warr. (A gathia). 
Helicopage cinerea, Prout. 


9,40 mm. Head bright green; lower half of face and 
fillet grey. Palpi 24, terminal joint 4; grey, basal half of 
under-surface whitish. Antennae grey. Thorax bright green 
with median and postmedian central grey spots. Abdomen 
pale grey with a dorsal series of large green spots; beneath 
whitish. Legs whitish ; anterior pair fuscous anteriorly. Fore- 
wings triangular, costa moderately arched, apex acute, termen 
strongly bowed, oblique; bright green with broadly suffused 
grey markings and strigulae; costal edge pale grey with 
darker strigulae; a rather large basal patch containing a 
fuscous subcostal spot and several green spots, towards dorsum 
this is darker, with a very oblique inwardly directed edge; 
succeeding this is a narrow irregular fascia connected with 
a transverse median bar, which runs into postmedian 
fascia; a very broad fascia with darker strigulae, its edges 
very irregular, extending on costa from 2 to apex, on dorsum — 
from 2 to tornus and adjacent part of termen, this forms an 
acute apical process, and contains a transverse sinuous line 
of fuscous dots at 3; a grey terminal line, cilia grey. Huind- 
wings with termen angled on vein 6, and more acutely so 
on vein 4; as forewings but with a small basal fascia only; 
postmedian fascia expanded into a large tornal blotch ex- 
tending from mid-dorsum to acute angle on termen, contain- 
ing a transverse series of fuscous dots and a dark wavy line 
from apex to tornus. Underside whitish; costa of forewings 
with large fuscous strigulae and a subapical blotch, from 
which arises a narrow transverse fascia; hindwings with a 
fuscous subterminal fascia thickest in middle. 

Unfortunately the male is unknown. In Helicopage the 
male antennae are pectinate, and the male frenulum abnorm- 


ally specialized. 


+ agree 
wn 
is 


279 


Northern Queensland: Kuranda, near Cairns, in January ; 
one specimen received from Mr. F. P. Dodd. Also from New 
Guinea. 

Gen. CyNEOTERPNA, Prout. 


Autanepsia, Turn., praeoce. 


Type C. wilson, Feld. 


Gen. HemicHiLoreis, Turn. 


HEMICHLOREIS THEATA, Turn. 
New South Wales: Taree. 


Gen. CrypsipHona, Meyyvr. 


In my revision I made C. melanosema the type of the 
genus. This was unfortunate, as Mr. Prout has pointed out, 
nor do I think it can be maintained. Although Mr. Meyrick 
did not specify the type, the name he has given to the genus 
(kpoydwvos, with hidden colour) clearly indicates that he 
intended occultaria as the type. 


CRYPSIPHONA EREMNOPIS, 0. sp. 
epepvwmes, dark. 
¢, 9,32 mm. Head brown-whitish irrorated with dark 


- fuscous. Palpi 2; fuscous, some brown-whitish scales on upper 


edge, base whitish beneath. Antennae fuscous; pectinations 
in male 5. Thorax fuscous mixed with brown-whitish. 
Abdomen grey. Legs, anterior pair dark fuscous [middle and 
posterior pairs broken off]. Forewings triangular, costa gently 
arched near base, thence nearly straight, apex obtusely 
pointed, termen bowed, oblique, crenulate; 11 anastomosing 
with 12 (1 male); brown-whitish suffused, and towards costa 
strigulated, with fuscous; markings fuscous; an indistinct, 
transverse, somewhat dentate line at 4; a transverse, linear, 
dark-fuscous, discal mark beneath mid-costa, surrounded by 
some brownish suffusion; a narrow fascia, ill-defined 


anteriorly, posteriorly sharply defined by whitish, at first bent 


outwards and very sharply dentate, abruptly bent inwards 
below middle, and ending as a fine line to ? dorsum; an 
indistinct, whitish, dentate, subterminal line, anteriorly 
edged by sharp fuscous teeth; some brownish suffusion 
between this and termen; a dark-fuscous terminal line; cilia 
fuscous, narrowly barred with white between veins. Hind- 
wings with termen rounded, crenulate; rather dark grey; 
an obscure, darker, dentate, postmedian line; a dark-fuscous 
terminal line; cilia as forewings. Underside whitish suffused 
with fuscous, with obscure dark postmedian line on both 
wings. 


280 


In the absence of the hindlegs I cannot be sure that this is 
a Crypsiphona, but the total absence of abdominal crests 
makes it probable. 

Western Australia: Cunderdin, in October, one male 
received from Mr. R. Illidge; Mount Barker, one female 
(L. J. Newman). 

Gen. Pineasa, Moore. 


Differs from T'erpna in having crests of scales on upper- 
surface of hindwings. The distinction seems natural and 
tenable. So far I agree with Prout, but cannot follow him 
in separating. from it a new genus Hypodoza; the former 
with cell of hindwings short, scale-tuft at its end; the latter 
with cell normal, scale-tuft before its end. I have carefully — 
noted (without actual measurement) the comparative length 
of the cell of the*hindwing in seven Australian species. The 
dorsal edge of the cell is longer than the costal, and I have 
made my comparisons from the length of the costal edge. 
In chlora it is about 2; in cinerea between 2 and 4; in 
emiliaria, muscosaria, myriosticta, and erebata about 4; in 
deteriorata about 2. These differences and slight variations 
in the position of the scale-tufts appear to me to be of specific 
value only. 

Type P. ruginaria, Gn., from India and Africa. 


PINGASA MUSCOSARIA, Gn. 


This species varies much according to locality. It would 
be easy to distinguish local races or subspecies, probably a 
longer series will show these to be connected by intermediate 
forms. . 
PINGASA ACUTANGULA, Warr. 


Q, 42-46 mm. Head brownish, on sides whitish. Palpi 
rather long, ascending ; terminal joint as long as:second joint, 
porrect; whitish. Antennae fuscous, towards base fuscous- 
whitish. Thorax whitish with a central brownish suffusion. 
Abdomen whitish suffused with fuscous and brownish; a 
double median reddish-brown line, enclosing crests, which 
are brownish; underside whitish. Legs, anterior pair 
fuscous, coxae whitish [middle and_ posterior pairs 
broken off]. Forewings triangular, costa gently arched, apex 
round-pointed, termen bowed, crenulate; whitish with fine 
pale-brown or grey irroration; lines fine, blackish, becoming 
reddish on dentations; first from } costa, acutely angled 
inwards beneath costa, then prolonged outwards nearly to 
middle of disc, where it forms a narrow quadrangular process, 
in which is included a brownish linear discal mark, return- 
ing it forms an acute angle on disc beneath subcostal angle, 


281 


beneath this a double phd eee te on vein | and ends on dorsum, 
| near base; second line from 3 costa towards termen, acutely 
dentate six times, then bent inwards to dorsum near middle, 
with a seventh dentation above dorsum ; terminal area darkly 
suffused with brown and fuscous beyond second line, and a 
short reddish line connecting sixth dentation with ‘tornus ; 
an obscure whitish dentate subterminal line; a suffused paler 
spot on termen below middle; a dark terminal line; cilia 
whitish obscurely barred with brownish. Hindwings similar 
| but without first line, discal mark small or absent. Under- 
side white; both wings with a blackish terminal band, and 
_ white apical and median terminal spots; forewings with linear 
_ discal mark. 
Easily recognized by the peculiarly angulated. first line 
of forewings. 
Northern Queensland: Coen River (W. D. Dodd), one 
| specimen in South Australian Museum; Kuranda (from F. P. 
Dodd in Coll. Lyell). Also from New Guinea. 


| PINGASA ATRISCRIPTA, Warr. 
Hypochroma mumta, Luc. 


| I do not know this species and have merely transcribed 
Prout’s identification. . 
| _ Northern Queensland: Cairns. Also from New Guinea. 


Gen. AEoLocHROMA, Prout. 
Type A. turneri, Luc. 


Mr. Prout refers here all the remaining Australian 
species of the group except paroptila (doubtfully) and 
percomptaria. These two he retains in Terpna, which he dis- 

| tinguishes by the frons being strongly protuberant. But in 

| percomptaria this is not the case, and being therefore doubtful 

of the validity of his distinction, I propose to retain all these 
species in Terpna except the type, defining the genus 
Aeolochroma by the simple male antennae. It differs from 
Actenochroma, Warr., in having strong abdominal crests. 


Gen. Terpna, H.-Sch. 

T. saturataria, Wlk., cannot be included in the Aus- 
tralian list at present. It may occur in Queensland, but Swin- 
hoe’s reference to Western Australia is almost certainly 
erroneous. 


—_ 


TERPNA UNITARIA, WI1k. (Tephrosia). 
Hypochroma acanthina, Meyr. 
I do not know this species. 


282 


TERPNA HYPOCHROMARIA, Gn. 


The male of this species has a small notch preceded by a 
small tuft of hairs on the dorsum of the antenna near its 
base. No doubt this is a scent-producing organ. 

Northern Queensland: Cape York. Queensland: Bris- 
bane, Toowoomba. New South Wales. 


Gen. STERICTOPSIS, Warr. 


Mr. Prout, who has examined the type of paratorna, 
Meyr., states (Gen. Ins. Hemith., p. 24) that it does not 
belong to’ this’ genus, for 10 is stalked with 7, 8, 9. It has 
scarcely any dorsal crests and the male antennal pectinations 
are short. Argyraspis, Low., is from the same locality pro- 
bably, and therefore may be identical with it. The two 
Gisborne examples, which I examined, agreed structurally 
with :nconsequens, Warr., which is from Duaringa, but I 
will not be sure that they are the same species. I accept, 
of course, Mr. Prout’s observations, but am unable for want 
of material to clear up the confusion, which at present un- 
doubtedly exists. 


ADDITIONAL LOCALITIES. 


Comostola laesaria, Wlk.—Q’land: Gayndah, Caloundra, Strad- 
broke Island, Mount Tambourine, Coolangatta, Rosewood, 
Toowoomba; N.S. Wales: Lismore. 

Pyrrhorhachis pyrrhogona, W1k.—Q’land: Gayndah, Rosewood. 

Chloéres citrolimbaria, Gn. —Q’land: Blackbutt, National Park 
(2-3,000' ft.).; N.S. Wales: Lismore, Port Hacking. 

Mixocera latilineata, Wlk.—Q’land: Gayndah, Caloundra, Too- 
woomba;, N.S. Wales : Lismore, Tabulam. 

Kuloxia meandraria, Gn.—N.S. Wales : Ebor, Mount Kosciusko 
(3,500-5,000 ft.). 

E. fugitivaria, Gn.—N.S. Wales: Glen Innes, Mount Kosciusko 
(5,000 ft.). ’ 

EK. pyropa, Meyr.—W. Austr.: Harvey. 

Chlorocoma cadmaria, Gn. —Q’land: Coolangatta; N.S. Wales: 

Glen Innes, 

. dichloraria, Gn.—Q’land: Brisbane, Blackbutt. 

. assimilis, Luc.—W. Austr. : Donnybrook. 

. externa, WIk. —Q’land : Toowoomba. 

. monocyma, Meyr.—S. Austr.: Port Augusta. 

. melocrossa, Meyr.—Q’land : Stradbroke Island. Coolangatta ; 

Tas. ? Hobart, Tasman Peninsula. 

Comibaena mariae, Luc.—Q’land: Gayndah, Rosewood, Too! 
woomba. 

Thalassodes veraria, Gn.—N. Terr.: Darwin; N.S. Wales: 
Lismore. 

Gelasma rhodocosma, Meyr.—N. Terr.: Darwin; N. Q’land: 
Cairns; Q’land: ’Gayndah. 

G. ocyptera, Meyr.—Q’land: Clermont, Gayndah, Toowoomba, 
Charleville. . 

G. albicosta, Wlk.—N. Terr.: Melville Island; N. Q’land: Cairns. 


eleleleke 


a) 


P| 


| 
| 
| 


283 4 


G. iosticta, Meyr.—N. Q’land: Herberton; Q’land: Stradbroke 
Island; N.S. Wales: Lismore. 
G. calaina, Turn.—Q’land: Montville (1,500 ft.) near Nambour, 
National Park (3,000 ft.), Toowoomba. 
G. centrophylla, Meyr. —N.S. Wales: Port Macquarie. 
G. floresaria, Wlk.—N. Q’land: Herberton. 
Hemithea insularia, Gn.—N. Terr.: Darwin. 
' Metallochlora decorata, Warr.—N. ‘Qland: Hlorbes tow. 
M. venusta, Warr. _N. Q’land: Atherton. 
Urolitha bipunctifera, Wlk.—Q’land: Gayndah, Moawaomba: N.S. 
Wales: Lismore. Also from Lord Howe Island. 
Uliocnemis partita, Wlk.—N. Q’land: Claudie River. 
Eucyclodes pieroides, Wi1k.—N. Terr.: Darwin; N. Q’land, Cook- 
town, Cairns; Q’land: Gayndah, Coolangatta ; N.S. Wales : 
Lismore. 
KE. fascinans, Luc.—N.S. Wales: Lismore. 
E. insperata, Wlk.—Q’land: Toowoomba; N.S. Wales: Lismore. 
E. metaspila, Wik.—Q’land: Nambour, Mount Tambourine. 
E. buprestaria, Gn.—Q’land : Coolangatta ; Tas. : Cygnet. 
Chlorodes boisduvalaria, Le G.—N.S. Wales : Ebor; Tas.: Hobart. 
Agathia, laetata, Fab. —Q’land: Nambour, Rosewood ; N. S. Wales : 
Lismore. 
| Crypsiphona occultaria, Don.—N. Terr.: Darwin; Q’land: Too- 
woomba, Charleville; N.S. Wales: Lismore; Vict.: Birchip; 
Tas.: Tasman Peninsula, Cygnet. 
Pingasa ‘muscosaria, Gn.—Q’land: Nambour, Toowoomba; N.S. 
Wales: Lismore, Ebor, Albyn River. 
. emiliaria, Gn.—N.S. Wales: Lismore. 
. myriosticta, Turn.—N.S. Wales: Lismore. 
. erebata, Wlk.—N. Terr. : Darwin; Q’land: Vepoocn: Cal- 
oundra. 
. chlora, Cram.—Q’land: Coolangatta. 
cinerea, Warr.—Q’land: Nambour, Caloundra, Toowoomba. 
Bina metarhodata, Wlk.—Q’land: Gayndah. 
T. hypochromaria, Gn.—Q’land: Gayndah, Nanango, Toowoomba ; 
N.S. Wales: Lismore. 
T. quadrilinea, Luc.—Q’land: eos N.S. Wale Lismore, 
Port Macquarie. 
T. percomptaria, Gn.—Q’land: Poogoomba. 
huma subaurata, Wlk.—N.S. Wales: Taree. 


a 


‘Heliomystis electrica, Meyr.—N.S. Wales: Mount Kosciusko 


. (5,000 ft.). | 
: Fam. BOARMIADAE. 


CLEORA LACTEATA, Warr. (Chogada). ‘| 


This name must be adopted for the species, which, follow- 
ing Meyrick, I have described under the name of wdlustraria, 


Wik. I have since examined the type of ilustraria and find 


that is referable to the species for which I have adopted the 


name acaciaria, Bdv. 


Also from New Guinea and New Britain. 


BoarRmia zascia, Meyr. 
Specimens from Armidale and Stanthorpe are much paler 


| than those from Victoria, the general coloration being ‘greyish, 
_ and the vertex of head is grey, but the face is always blackish. 


12 


284 


Queensland : Stanthorpe, in October. New South Wales. 
Armidale. Victoria: Melbourne, Beaconsfield. 


BoaRMIA PANCONITA, Turn. 


Nearly allied to B. zascia. It is darker than the northern 
examples of this species, from which it may be always dis- 
tinguished by the lower part of the face being white, and by 
the crescentic discal mark on the hindwing. [The female 
example with wholly blackish face, which I formerly referred 
to this species, is an example of zascia.]| The Gayndah 
examples apparently represent a distinct local race. 

Queensland: Gayndah, Stanthorpe, in October. 


_  BOARMIA DESTINATARIA, Gn. 

Also allied to the two preceding species, and like them: 
variable, but readily ,distinguished by the paler suffused 
coloration more or less tinged with ochreous, and the absence 
of any black on the face. / & 

Queensland : Stanthorpe, in October. New South Wales: 
Ebor, Sydney, Katoomba. Tasmania. 


BOARMIA PISSINOPA, 0. sp. 
muowwros, black as pitch. 


3,42mm. Head, palpi, antennae, and thorax blackish. 
Antennal pectinations in male 10, apical ¢ simple. 
Abdomen on dorsum fuscous becoming blackish towards base; 
lower-surface, sides, and tuft grey-whitish. Legs fuscous; 
posterior pair grey. Forewings triangular, costa nearly 
straight, apex round-pointed, termen bowed, oblique, slightly 
crenulate; blackish; markings intensely black; a fine trans- 
verse line from 4 ae bent strongly inwards beneath costa, 
and again bent to % dorsum ; a thicker oblique shade from 
mid-costa to dorsum before midule ; a transverse, median, 
subcostal discal mark; a slightly dentate line from 3 costa, 
strongly bent inwards to mid-dorsum; a faint, incomplete, 
dentate subterminal line; a fine terminal line; cilia dark 
fuscous. Huindwings with termen gently rounded, obtusely — 
dentate; as forewings but without first line, other lines trans- 
verse, gently rounded. 

In colour this species resembles Melanodes anthracitaria, 
Gn., and ‘both are adapted for concealment on tree-trunks 
blackened by fire. 


Western Australia: Perth, in October; one specimen. 


BoaRMIA MACULATA, Luc. 


Queensland: National Park (3,000 ft.), in March; a 
series taken at light. These agree with two examples from — 


>] 


285 


Kuranda which I have identified at maculata, Luc., in 
structure of male antennae, neuration (10 and 11 stalked, 
free; 6 males and 4 females), and markings, but they are 
larger (52-58 mm.) and much greener in coloration. 


ABRAXAS SPOROCROSSA, Nn. sp. 
o7TOpoKpog aos, with spotted border. 


3,9, 46-50 mm. Head yellow with three fuscous dots 
on crown and sometimes another on face. Palpi fuscous, 
towards base yellowish. Antennae fuscous; ciliations in male 
4. Thorax fuscous; middle of patagia and two posterior 
dots yellow. Abdomen fuscous on dorsum; bases of segments 
broadly yellow, each yellow bar containing a pair of lateral 
spots ; ventral surface yellow with paired fuscous spots. Legs 
fuscous-grey ; coxae and posterior femora partly yellowish. 
Forewings triangular, costa strongly arched, apex rounded, 
termen bowed, oblique; blackish; a yellow dot beneath costa 
near base, followed by a median whitish dot, which is some- 
times connected with a subcostal dot at 4, these are more 
or less yellow tinged; a quadrangular white spot beneath 
4+ costa; a triangular blotch on mid-dorsum, its apex acute 
and reaching nearly to middle of disc; a white blotch beneath 
2 costa, irregular in outline, reaching below middle of disc, 
convex posteriorly, concave and more or less wavy anteriorly, 
followed by a minute subcostal dot; a white dot before tornus, 
sometimes prolonged into disc; a subterminal series of six 
or seven small quadrangular white spots, the two central 
reduced to dots; cilia blackish. Hindwings with termen 
gently rounded; white; a triangular basal blackish blotch 
to +; a blackish terminal band containing a series of quad- 
rangular white dots; cilia blackish. Underside similar. 

Northern Queensland: Claudie River, in December; two 
specimens taken by Mr. J. A. Kershaw. Type in National 
Museum, Melbourne. 


Gen. XYLODRYAS, n. gen. 
EvAodpvas, a Woodnymph. 


Frons flat. Tongue well developed. Palpi moderate, 
porrect; basa] and second joints shortly rough-scaled ; 
terminal joint short. Antennae in male simple, minutely 
ciliated. Thorax with a small posterior crest; slightly hairy 
beneath. Abdomen not crested. Femora smooth. Posterior 
tibiae in male not dilated. Forewings broadly triangular, 
costa strongly arched towards base, termen excavated between 
veins 4 and 6; in male without fovea; 2 from 2, 7, 8, 9, 10 
stalked, 10 connected with 8, 9 beyond 7, 11 connected with 


286 


12. Hindwings obtusely angled on veins 4 and 7; 2 from , 
3 and 4 widely separate, 6 and 7 separate, 8 closely approxim- 
ated to cell to beyond middle. 

Type X. leptoxantha, which I formerly included, while 
pointing out the differences, with Coelocrossa, Turn. On 
reconsideration it appears to me generically distinct, and per- 
haps not closely allied. Apart from minor differences the 
structure of vein 8 of hindwings affords an important dis- 
tinction. I suspect some affinity with Lyellana, Turn., and 
Lophosema, Turn. 

I think this is probably, with a few other Geometridae, 
part of the aboriginal fauna of the Eastern Islands before 
they became part of the Australian continent. _ 


XYLODRYAS LEPTOXANTHA, Turn. 

I took one male on the wing by lantern light in the 
National Park, Queensland (2,500-3,000 ft.), in December. 

The species is not confined to the mountains, for I have 
received from Mr. G. N. Newman a very similar specimen 
taken at Rous, near Lismore, New South Wales. A second 
example taken in the National Park in March is a very dis- 
tinct aberration, purplish-grey, with faint lines, little irrora- 
tion, but a small whitish spot near base of forewing, and 
others near termen of both wings. 


BURSADA FLAVANNULATA, Warr. 


3, 9, 24-30 mm. Head and thorax blackish; face and 
palpi ochreous-whitish or grey-whitish. Antennae blackish; 
pectinations in male 12, in female 4. Abdomen blackish; a 
transverse subbasal yellow or orange line on dorsum. Legs 
fuscous. Forewings triangular, rather narrow, costa gently 
arched, apex rounded, termen bowed, oblique; blackish; an 
oblique oval yellow or orange blotch extending from beneath 
2 costa to above termen beyond tornus; cilia blackish. Huind- 
wings with termen rounded; yellow or orange; a blackish 
terminal band, sharply defined, broad at apex and tornus, 
narrower on mid-termen, ending rectangularly above tornus, 
but giving off a. subdorsal streak towards base; cilia blackish. 
Underside similar. 

Northern Queensland: Claudie River, in March; two 
specimens taken by Mr. J. A. Kershaw. Also from New 
Guinea. 


Gen. CLEPSIPHRON, n. gen. 
krAefippwv, deceiving. 
Frons flat. Tongue present. Palpi short, porrect, pro- 
jecting only slightly beyond frons; second joint shortly rough- 
scaled; terminal joint very short, depressed. Antennae in 


287 


male simple, minutely ciliated. Thorax and abdomen without 
crests; thorax smooth beneath. Femora smooth; all tibial 
| spurs present; inner twice as long as outer. Forewings with 
base of costa rounded; in male without fovea; 2, 3, 4 equi- 
distant, 5 from middle of cell, 6 from upper angle, 7, 8, 9, 
10, 11 stalked from considerably before angle, 11 only short- 
stalked, connected first with 12 and then with stalk of 7, 8, 
9, 10. Hindwings broad; cell about 2; 5 absent, 6 and 7 
separate, the latter arising from shortly before angle, 8 con- 
nected with cell near base, thence diverging. 

A peculiar genus, but probably related to Peridelias, 
Turn., Aplochlora, Warr., and Parametrodes, Warr. 


CLEPSIPHRON CALYCOPIS, N. sp. 

KaAvkwris, roseate. 
3, 20mm. Head ochreous-grey ; face with some reddish 
scales; posterior margin of eyes reddish. Palpi ochreous- 
whitish; second joint barred with reddish in middle and at 
apex. Antennae whitish-grey. Thorax purplish-grey. 
Abdomen reddish-grey ; tuft ochreous-whitish. Legs ochreous- 
whitish; anterior femora and tibiae reddish tinged; anterior 
tarsi fuscous tinged. Forewings broadly triangular, costa 
strongly rounded at base, thence slightly arched, apex rect- 
angular, costa not oblique, slightly sinuate; purple-fuscous ; 
base of costa purple; an ill-defined darker basal patch; an 
outwardly curved fuscous line from 4 costa to dorsum before 
middle, indistinct towards costa, towards dorsum well defined 
and mixed with orange; a line from #% costa, at first out- 
wardly curved, but bent inwards and then angled outwards 
above dorsum, ending on dorsum before tornus, orange be- 
coming fuscous towards costa; termen with a narrow, 
irregularly-indented, yellow margin; cilia pale yellow. Hind- 
wings with termen wavy and slightly angled on vein 4; 
purple-fuscous, the greater part of disc suffused with reddish 
and orange with small purple-fuscous strigulae; terminal 
_Margin and cilia as forewings. Underside grey with traces of 
whitish postmedian line, and with whitish terminal margin. 

Northern Queensland: Evelyn Scrub, near Herberton, 
in January; one specimen received from Mr. F. P. Dodd. 
Type in Coll, Lyell. 


Gen. PIcROPHYLLA, n. gen. 
miKpopvaAdros, with pointed wings. 


Frons with an anterior tuft of scales. Tongue well 
~ developed. Palpi rather short, porrect; second joint rough- 
haired; terminal joint short. Antennae of male simple, 


288 


ciliations minute. Thorax and abdomen without crests; thorax 
slightly hairy beneath. Femora smooth; posterior femora of 
male dilated with internal groove and tuft. Forewings in 
male without fovea; 10 and 11 long-stalked, 10 anastomosing 
with 8, 9 beyond 7. Hindwings with apex produced to a 
sharp point on vein 7; 3 and 4 approximated at origin; 6 and 
7 separate, 7 arising before angle of cell, 8 closely ap- 
proximated to cell for nearly its whole length. : 
Probably allied to T'essarotis, Warr., which approaches 
it closely in wing shape, but has 10 and 11 arising separately. 


PICROPHYLLA HYLEORA, Nl. sp. 
vAnwpos, of the woods. 


3, 9,40 mm. Head fuscous-brown. Palpi 14; fuscous- 
brown. Antennae ochreous-whitish, dorsum except towards 
apex suffused with fuscous-brown. Thorax brown-whitish; a 
postmedian pair of fuscous dots. Abdomen brown-whitish ; 
paired fuscous dots on dorsum of second and third segments. 
Legs whitish-ochreous speckled with dark fuscous. Forewings 
triangular, costa slightly arched, apex acute, produced, 
termen sinuate beneath apex, angled on vein 4, thence slightly 
concave to tornus; brown-whitish with sparsely scattered, 
dark-fuscous, transverse strigulae, more numerous on costa, 
towards base, and towards termen; a suffused fuscous line 
from 4 costa with two posterior teeth, beneath costa and in 
middle, obsolete towards dorsum; a fine, straight, fuscous- 
brown line from costa before apex to 4 dorsum, succeeded by 
a parallel row of fuscous dots; a dark-fuscous discal dot 
beneath 2 costa; cilia fuscous, on costa and from beneath 
apex to angle brown-whitish. Hindwings produced to a 
sharp point on vein 7, termen beneath this sinuate, thence 
nearly straight; as forewings with fewer strigulae; without 
first line; second line median ; a subterminal series of fuscous 
dots; cilia brown-whitish. Underside similar. 


Queensland: Eumundi, near Nambour, in January; 
National Park (3,000 ft.), in March; two specimens. 


CASBIA RHODOPTILA, Turn, 


In addition to the type I have now a female (26 mm.) 
from Northern Territory, Darwin (G. F. Hill) without spots 
on forewing; and a male (30 mm.) from Queensland, Strad- 
broke Island, in August, with discal dot, but without pos- 
terior spot. The reddish head and tegulae form a good 
distinguishing mark of this species. In all my three examples 
vein 11 of forewing anastomoses with 12. 


289 


IDIODES ARGILLINA, N. sp. 
apyAdwos, clay-coloured. 


3, 44 mm. Head and thorax brown. Palpi about 1; 
brown. Antennae dark grey. Abdomen grey; dorsum brown 
towards base. Legs grey; anterior pair fuscous. Forewings 
broadly triangular, costa gently arched, apex obtusely pointed, 
termen slightly bowed, slightly oblique; brown with numerous 
fine transverse fuscous strigulae, these are most numerous on 
costa, present also towards margins, and across main veins; 
a large suffused fuscous blotch, its margins composed of 
coalesced strigulae, extends on costa from middle to apex, 
narrowing dorsally it terminates abruptly on vein 2; an in- 
distinct, very narrow, interrupted, pale, oblique line from 
apex, traversing the dark blotch towards # dorsum; cilia 
brown. Hindwings with termen slightly rounded; colour and 
strigulae as forewings, but without blotch; a suffused darker- 
brown line from mid-dorsum towards 4 costa; in this a small 
fuscous discal spot; cilia brown. Underside similar. 

Nearest J. ficteélis, Turn. 


Queensland: National Park (3,000-3,500 ft.), in 
January; one specimen. 


Gen XeENomusA, Meyr. 


Frons smooth, not projecting. Tongue well developed. 
Palpi short (1 or less), hairy beneath. Antennae in male 
simple or bipectinate. Thorax not crested; beneath hairy. 
Abdomen without crests. Femora smooth-scaled. Posterior 
tibiae with all spurs present; in male not dilated. Forewings 
with apex uncinate and slightly produced; cell over 4, dis- 
cocellulars nearly straight, or inwardly curved, 2 from 3, 
3 and 4 separate, 5 from or from above middle, rather weakly 
developed, 6 separate or short-stalked, 10 from cell or short- 
stalked wut 7, 8, 9, 10 and 11 free. Hindwings with 2 


from 2 or 3, 3 and 4 separate, 5 obsolete or weakly developed, 


6 and 7 separate, 12 closely approximated to cell as far as 
middle. 


Meyrick placed this among the Oenochromidae. In X. 
metallica, vein 5 of hindwings is obsolete, being concealed in 


a fold of the wing membrane; in X. rubra it is present, but 


weak. I think the two must be regarded as congeneric in 


spite of this and the difference in antennal structure. X. 


monoda, the type species, I have seen, but have no specimens 
for examination. The genus should be placed, I think, in 
Boarmiadae, of which it is a primitive form. In X. rubra 
a forked median vein is plainly visible in the cell. 


290 


XENOMUSA METALLICA, Luc. 

3, 34 mm.; 9, 40-45 mm. Head brownish or grey; 
two whitish spots or a white line on lower edge of face. 
Palpi in male 4, in female #; whitish or whitish-ochreous, 
apex blackish. Antennae grey; in male simple, minutely 


ciliated. Thorax brownish or grey. Abdomen brownish or — 


grey with sparsely scattered blackish scales. Legs ochreous- 
whitish ; tibiae and tarsi annulated with dark fuscous. Fore- 
wings elongate-triangular, narrower in male, costa bisinuate, 
more strongly so in male, apex uncinate, produced, termen 
bowed, oblique; 10 short-stalked (1 male and 7 females) ; 
brownish or grey usually with sparsely scattered blackish 
scales; usually a whitish-ochreous spot on base of costa; a 
fuscous or brownish line from 4 costa very obliquely ‘out- 
wards, sharply angled beneath costa, thence very obliquely 
inwards to dorsum near base; a similar line, posteriorly edged 
with whitish, from beneath costa before apex, nearly straight, 
to dorsum before middle; usually a minute, blackish, median, 
discal dot beneath costa; apex fuscous preceded by whitish ; 
a short oblique line or fuscous shade from apex to beneath 
second line; cilia fuscous. Hindwings with termen very 
slightly rounded, tornus prominent; colour and cilia as fore- 
wings; a straight transverse brownish or fuscous line at 4; 
a white, median, discal dot. 

Northern Queensland: Kuranda, in April; one male. 
Queensland: Montville, near Nambour, in March; Brisbane, 
in January and March; seven females. 


XENOMUSA RUBRA, Luc. 

2, 50 mm. Head pale reddish; face reddish-orange. 
Palpi bs; Podge Puree. Antennae reddish-orange; in female 
shortly bipectinate (14), apical 4 simple. Thorax pale reddish. 
Abdomen ochreous. Legs ae ochreous. Forewings tri- 
angular, costa gently bisinuate, apex produced, slightly 
uncinate, termen sinuate, oblique; 10 from cell; reddish- 
orange without markings; cilia reddish-orange. Hindwings 
with termen slightly rounded, tornus rather prominent; as 
forewings. 

My description is taken from Dr. Lucas’ type, which is 
in my possession, and still, I believe, remains unique. 

Queensland: Brisbane. 


Gen. Dirce, Prout. 

Oenone, Meyr., pracocc. 
; This genus must be transferred to the Boarmiadae, for 
descaling shows that vein 5 of the hindwings is absent. ‘Pre- 
vious authors have been deceived by the presence of a 


291 


persistent fold of the wing-membrane in the normal position of 
this vein. On the other hand, Diceratucha, Swin., has vein 5 
of hindwings sufficiently well developed, and must be retained 
in the Oenochromidae. The two genera agree in the neura- 


| tion of the forewing, in which the areole is of a primitive 


form, and no doubt there is real relationship between them. 
In fact, the latter genus is probably very near the point, 
where the primitive stem of the Boarmiadae diverged from 
the Oenochromidae. 

I can see no valid grounds for the conjectures of Meyrick 
and Prout for any near relationship to Brephos, which has 
completely lost the areole. Its points of resemblance to 
Dirce are merely superficial (general hairiness and colour 
scheme) and adaptational. MHairiness is a common character 
in genera of mountain localities, and is probably a protection 
against the dampness of mountain mists. 


_DrIRcE AESIODORA, 0. sp. 
ductodwpos,a fortunate gift. 


36,9, 26-30 mm. Head blackish with a white central 
spot on crown; face white, hairs on margins blackish. Palpi 
projecting somewhat beyond frons; white; some hairs, apex of 
second joint, and whole of terminal joint blackish. Antennae 
blackish; in male thickened, serrate, and minutely ciliated. 
Thorax blackish irrorated with whitish. Abdomen dark fus- 
cous; irroration, apices of segments, and some hairs in tuft 
ochreous-whitish. Legs blackish; tibiae and tarsi annulated 
with white; posterior pair whitish on posterior surface. Fore- 
wings triangular, costa arched near base, thence slightly 
sinuate, apex rectangular, termen slightly bowed, not oblique; 
blackish mixed with grey and white; markings white; a basal 
spot; a bar from costa near base uniting with another from 
costa at 4+, to form a fascia, which extends on dorsum from 
near base to 4, and is sharply toothed posteriorly above 
dorsum ; two suffused spots on dorsum before and after middle, 
the first larger and produced across disc towards costa; a spot 
- on mid-costa; a narrow fascia from # costa to ? dorsum, pos- 
teriorly suffused, anteriorly sharply defined, with a circular 
anterior process containing a central blackish dot beneath 
costa; a slender, interrupted, subterminal line; a series’ of 
wedge-shaped black marks beyond this, separated in female 
by some whitish suffusion ; terminal edge blackish ; cilia blackish 
barred with white. Hindwings with termen rounded; 
blackish with a large central orange blotch, sometimes preceded 
by a small triangular spot near base; cilia orange barred with 
blackish, on apex and costa blackish. Underside pale orange ; 
forewings with basal patch, oblique median fascia, costal spot 


292 


and terminal fascia blackish; hindwings with oblique fascia 
from + costa to mid-dorsum, and broad band from costa before 
middle around apex and termen to tornus. 

Tasmania: Cradle Mountain (3,000-3,500 ft.), in January ; 
four specimens received from Dr. R. J. Tillyard. 


Fam. OENOCHROMIDAE. 


OENOCHROMA LISSOSCIA, N. sp. 
Aurcooxos, smoothly shaded. 


Q, 46-48 mm. Head, palpi, and thorax grey. Antennae 
dark grey. Abdomen grey with a few blackish scales; under- 
surface reddish. Legs grey, partly reddish tinged ; tarsi 
fuscous. Forewings elongate-triangular, costa bisinuate, apex 
acute, termen strongly bowed, becoming straight towards 
tornus; grey with a few scattered blackish scales; some fine 
fuscous-brown transverse strigulae from basal half of costa; a 
fuscous-brown suffusion on costa from middle nearly to apex, 
leaving costal edge for a short distance at about # whitish; 
a fine blackish line from costa shortly before apex to 2 dorsum, 
outwardly bowed in middle, towards dorsum preceded by a 
fuscous-brown parallel line, costal half edged posteriorly by 
whitish, which extends to apex; some grey-brown suffusion 
on termen, preceded in middle by a suffused blackish spot; 
cilia fuscous-brown. Hindwings with termen slightly rounded, 
tornus prominent, rectangular; as forewings but with blackish 
line antemedian, straight, preceded by a fuscous-brown line, 
which diverges somewhat towards costa; no subterminal spot. 
Underside similar; but forewings with a blackish spot on 
costa near apex, with two blackish dots on veins beneath it, 
and no brownish suffusion ; disc purplish tinged with darker 
median transverse line; hindwings with a purplish antemedian 
fascia; posteriorly to this brownish, with suffused reddish 
subterminal spot between veins 3 and 4. : 

Exceptional in the genus is that veins 10 and 11 of 
forewings arise separately from the cell. 


Queensland : National Park (3,000 ft.), in March; three - 


specimens taken at light. 


OENOCHROMA ARTIA, 0. Sp. 

dptios, perfect. 

g, 38 mm. Crown of head yellow with a dark-reddish 
anterior line; face whitish. Palpi whitish with a few crimson 
scales. Antennae brownish-ochreous; pectinations in male 1}. 
Thorax pale green; bases of patagia yellow; pectus whitish, 
margin of eyes and forewings ochreous-yellow. Abdomen 
whitish. Legs whitish irrorated with crimson. Forewings 


pe 3 


293 

triangular, costa straight, apex pointed, termen nearly 
straight, oblique; pale green; a yellow line along 
costa to #; an oblique yellow line from mid-dorsum, 
moderately broad, but narrowing to extremity, which hes just 
beneath 2 costa; terminal edge whitish; cilia pale yellow. 
Hindwings with termen rounded; whitish; a yellowish suf- 
| fusion on mid-dorsum giving rise to a short transverse line ; 
a greenish suffusion on dorsum before tornus; a large round 
brownish-ochreous subtornal blotch; cilia whitish, around 
tornus yellow. Underside of forewings similar to upperside, 
but paler and without oblique line; of hindwings greenish- 
white, tornal blotch anteriorly orange, posteriorly deep 
_ crimson. 

Western Australia: Dardanup, in October; one specimen 
_ received from Mr. G. F. Berthoud. Type in Coll. Lyell. 


Gen. Noreia, WIk. 
NoREIA LOXxosTICHA, Turn. (Idiodes). 


| I have since received a male example, which shows a 
® small hairy tuft on underside of hindwing over vein 2, and 
_ has the posterior tibiae dilated with internal groove and tuft. 
The species has some close allies in the Indo-Malayan region, 
and I will not be sure of its distinctness. 
Northern Queensland: Kuranda in April and May; two 
specimens received from Mr. F.-P. Dodd. 


Gen. CELERENA, Walk. 


Face smooth. Tongue well developed. Palpi moderate, 
porrect; second joint shortly rough-haired; terminal joint 
short, with smoothly adpressed hairs. Antennae rather more 
than 4; in male shortly ciliated, usually with a small tuft of 
scales about middle, beyond this with moderately long bristles. 
Thorax densely hairy beneath, usually with an expansile 
posterior tuft of hairs. Abdomen of male usually with a 
basal tuft of long hairs on under-surface. Femora densely 
hairy. Posterior tibiae of male dilated with inner expansile 
tuft of hairs, long crooked median spurs, inner terminal 
spur only, its apex prolonged into a strong outer horny 
process. Forewings in male with a deep basal furrow beneath 
in cell: 7, 8, 9 stalked, 10 and 11 stalked, their stalk anas- 
| tomosing strongly with 12, 10 connected with 8, 9. Hind- 
wings with 5 from above middle of cell, 6 and 7 separate, 
8 moderately remote from cell, connected with it by an 
_ oblique bar near base. 

Type C. divisa, Wlk. An Indo-Malayan genus which is 


§ rather largely represented in New Guinea. 


294 } 


CELERENA GRISEOFUSA, Warr. 


3, 52 mm. Head yellow. Palpi yellow; apex of 


terminal joint fuscous. Antennae fuscous; in male minutel 


ciliated, apical 4 with moderately long bristles (14). Thorax . 
grey, anteriorly suffused with ochreous. Abdomen grey, sides | 


and under-surface ochreous. Legs grey; coxae and under- 
surface of posterior tibiae pale ochreous; first joint of pos- 


terior tibiae with an internal hairy tuft. Forewings tri- — 


angular, costa straight to #, thence arched, apex round- 
pointed, termen straight, oblique; grey with some yellow 
suffusion, most marked in costal half of cell; an incomplete 
narrow yellow fascia from 2 costa, outwardly oblique, inter- 
rupted in middle, then curved slightly inwards, and not 
reaching tornus; a band of yellow suffusion posterior and 
parallel to this; cilia grey. Hindwings with termen gently 
rounded; yellow; a moderate grey terminal band edged 
anteriorly by a blackish line and suffusedly prolonged along 
dorsum for some distance; cilia grey. Underside of forewings 
dark fuscous with a moderate yellow postmedian fascia not 
reaching: tornus; of hindwings yellow with a dark-fuscous 
terminal band. 

Northern Queensland: Claudie River, in March; one 
specimen taken by Mr. J. A. Kershaw. Also from New 
Guinea (Fergusson Island). 


Fin 


pine 


295 


THE FLORA AND FAUNA OF NUYT’S ARCHIPELAGO AND 
THE INVESTIGATOR GROUP. 
NO. 4-COLEOPTERA. 


By Artuvr M. Lra, F.E.S., Museum Entomologist. 
Contribution from the South, Australian Museum. 


[Read September 14, 1922. ] 
Puate XIII. 


The small but interesting collection of Coleoptera here 
dealt with was obtained on the islands by Prof. F. Wood 
Jones, and presented to the South Australian Museum. As 
_ he was specially interested in the mammals, and had but a 
short time on each island, the time available to colléct insects 
was always small, and those obtained are mostly sand- 
frequenting species, taken on or near beaches, and usually 
of wide distribution in Australia; even the new species, at 
present known only from the islands, will probably be 
eventually found on the mainland. Some of the Tene- 
brionidae were sent to Mr. H. J. Carter, for his opinion, 
and his descriptions of two new species are incorporated. 


CARABIDAE. 


Ectroma benefica, Newm. Numerous specimens of a 
pale variety of this species were obtained in rats’ nests on 
Franklin Island. 

Scopodes sigillatus, Germ. Six unusually small speci- 
mens were taken on Franklin Island. 

Lecanomerus flavocinctus, Blackb. Flinders Island. 


STAPHYLINIDAE. 


Hyperomma lacertinum, Fvl. This curious wingless 
species was previously known only from King George Sound. 
Prof. Wood Jones took one specimen on Franklin Island and 
Sir J. C. Verco another on St. Francis Island. 


SCYDMAENIDAE. 
Scydmaenus franklinensis, n. sp. 

. 3d. Bright castaneous, palpi and tarsi paler. Head and 

prothorax (except in middle) with fairly long and somewhat 

_ golden, or pale reddish hairs, similar but sparser hairs on 

elytra, but fairly numerous about base; under-surface with 
short pubescence. 

Head rather small; with sparse and small, but (when not 

concealed by clothing) sharply defined punctures. Eyes small 


296 


and prominent. Antennae rather long and thin; club four- 
jointed, its first joint scarcely longer than the preceding one 
but distinctly wider, apical joint almost as long as two pre- 


ceding combined. Prothorax moderately long, front parts : 


gently convex, flattened about base, each side of base with 
a transverse semidouble fovea; with minute scattered punc- 
tures. Elytra subovate, widest just before the middle, where 
they are about twice the width of prothorax, a fairly large 
impression on each side of base; with sparse, indistinct punc- 
tures. Subapical segment of abdomen incurved in middle of 
apex, the incurvature bounded on each side by a slight pro- 
jection. Front femora stout, the middle and hind ones 
pedunculate, front trochanters dentate. Length, 1°25-1°5 mm. 

@. Differs in having antennae shorter, elytra shorter 
and wider, abdomen simple, front trochanters unarmed, and 
front tibiae thinner and less curved at the tip. 


Hab.—South Australia: Franklin Island (Prof. F. Wood 
Jones). Type, I. 15360. . 

Almost the exact size of S. parramattensis, but more 
uniformly coloured, clothing different and club thinner; 
about the length of S. brevipilis, but narrower, club thinner 
and elytral clothing different. Of the species previously 
known from South Australia, S. depressus is much smaller, 
with wider elytra, shorter antennae, etc.; S. griffith: and S. 


fuscipalpis are much smaller, narrower, and darker, etc., and: 


S. impavidus has wider and glabrous elytra, etc. From 
some directions the hairs appear to form a loose fascicle on 
each side at the base of the head. When viewed at a right 
angle the armature of the male abdomen is inconspicuous, 
but when viewed from in front the projections appear as small 
subconical tubercles. 
DERMESTIDAE. 
Dermestes cadaverinus, Fab. Franklin Island. 
D. vidginus, Fab. Franklin Island. 


SCARABAEIDAE. 


Pimelopus dubius, Blackb. Franklin Island. 
P. porcellus, Er. Flinders Island. 


TENEBRIONIDAE. 
- Saragus posidonius Carter, n. sp 


Oval, convex, nitid black, oral organs, antennae and tarsi 
castaneous. 


Head finely punctate, antennae with joint 3 half as long 
again as 4, 8-11 as wide as long; epistoma a little incurved in 


7 
} 
i 
} | 
} 


297 


front. Prothorax moderately convex, subtruncate at apex be- 
tween the widely rounded anterior angles, foliate margins wide, 
sides arcuately diverging from apex to base, posterior angles 
produced and falcate; disc minutely punctate, the foliation 
concave with a strongly recurved border. Elytra almost as 
wide as long (9x8 mm.), convex, horizontal margin moder- 
ately wide at base, narrowing at apex; irregularly, coarsely 
substriate- _punctate, both rows, and punctures in rows closely 
placed, the punctures smaller and sometimes discontinuous 
near suture, larger and more regular towards sides, each 
4 rows bounded by a costate interval, with a less raised 
and more irregular costa half-way between each of these— 
the suture also costate—a lateral row of larger punctures, 
the explanate margins slightly wrinkled. Prosternum and 
episterna finely pustulose, abdomen striolate. Legs moder- 
ately long, tibiae with margins entire, terminal spines short, 
fore tarsi with basal joints wide. Dimensions, 12x 8 mm. 

Hab.—Neptune Island. 

Two examples show a species nearest to S. carinatus, 
Breme, but of smaller size and stronger sculpture. In con- 
vexity and style of sculpture it is suggestive of S. brunnipes, 
Boisd., but the punctures are coarser, the costae more pro- 
nounced, and the foliation of pronotum and elytra wider than 
in that species. The name suggests its habitat. Type, 
I. 15356. 


Saragus oleatus Carter, n. sp. 
Pl. xiii., fig. 1. 

Widely oval, convex, brilliantly nitid black, oral organs, 
antennae and tarsi castaneous. 

Head minutely, sparsely punctate, epistoma truncate, 
antennae with joint 3 proportionately shorter than in 
posidonius. Prothorax very convex and mirror like, apex 
narrowly arcuate, the anterior angles more squarely rounded, 
the posterior more acute, the foliate margins narrower and 
more deeply hollowed, the. sides less strongly arched, the 
recurved border considerably thicker than in the preceding 
species; disc submicroscopically punctate. Elytra nearly as 
wide as long (8x7; mm.), very convex, lateral margin nar- 
rower than in the preceding ; coarsely and unevenly striate- 
punctate, the 4 sutural rows of large punctures on each 
tending to confluence, rows 5 and 6, also 7 and 8, delimited 
by three costate intervals ; beyond these the seriate punctures 
“uneven in size, the intervals irregularly convex, the suture 
carinate throughout ; a lateral row of large punctures. Pro- 
sternum finely pustulose at sides, abdomen striolate. Legs 
shorter than in S. posidonius. Dimensions, 11x 74 mm. 

Hab.—Pearson Island. Type, I. 15357. 


298 


I have examined three examples of this species, which 
is more closely allied to S. brunnipes, Boi., then the pre- - 
ceding, but with a similar style of sculpture. It is remark- 
able for the apparently highly varnished surface, its polished 
and convex pronotum, coarsely punctate elytra with its 
irregular series and costate intervals. Wider and more convex 
than S. brunnipes; it is narrower and less convex than S. 
sphaeroides and S. french. 

Saragus brunnipes, Boi. Four specimens from South 
Neptune Island represent a rather coarsely punctate variety 
of this species. The species was also taken on Black Rocks. 

Pterohelaeus simplicicollis, Blackb. One specimen from 
Franklin Island, and another from St. Francis Island, iden- 
tified by Mr. Carter as probably belonging to this species. 

. PB. mtidissemus, Pasc. A single specimen from Flinders 
Island noted by Mr. Carter as having seriate punctures on 
elytra a little larger than on the typical form. 

P. ovalis, Blackb. St. Francis Island. 

Helaeus modicus,. Blackb. A very interesting series of 
33 specimens was taken on Franklin Island, ranging in length 
from 18 to 25 mm. Of these 14 have the curved portion on 
the left of the apex of the thorax on top of the right portion, 
and 16 have the right on top of the left; the difference is 
not sexual; on three the curved parts do not touch, being 
separated about half a millimetre. The species was also taken 
on: Goat Island (pl. xii., fig. 2). 

H. castor, Pasc. Franklin Island. 

Brises duboulayi, Bates. Franklin Island. 

Micrectyche nana, Pasc. A specimen from Franklin 
Island, identified by Mr. Carter as probably belonging to this 
species. . 

Caediomorpha heteromera, King. Black Rocks, St. 
Francis, Flinders, and Franklin Islands. 

Hyocis bakewelli, Pasc., var. pallida, Macl. St. Francis 
Island. 

Trachyscelis ciliaris, Champ. Franklin, Hyre, and Flin- 
ders Islands. 

Cestrinus aspersus, Blackb. Franklin Island. 


ANTHICIDAE. 


Anthicus strigosus, n. sp. 
Pl xt. 3. 

Head and prothorax dark reddish-brown, elytra almost 
black, legs, antennae and palpi more or less reddish, tarsi 
paler. Elytra moderately clothed with pale, subdepressed 
pubescence. 


299 


Head moderately large, parallel-sided for a short distance 
behind eyes, and then hind angles rather strongly rounded ; 
with crowded and small punctures, many of which are longi- 
tudinally confluent; with a narrow and continuous shining 
median line. Eyes small, medio-lateral and very prominent. 
Antennae rather long. Prothorax very little longer than 
wide, sides strongly rounded, but suddenly narrowed near 
base; densely and finely longitudinally strigose. Elytra 
elongate-elliptic, shoulders completely rounded off; with not 
very dense and rather small, but sharply defined punctures, 
becoming very small posteriorly. Legs moderately long. 
_ Length, 2-2°25 mm. 


Hab.—South Australia: Port Lincoln (Rev. T. Black- 
burn), Eyre Island (Prof. F. Wood Jones). Type, I. 15278. 


The prothorax is deeply striated and the head has a 
shining median line as in A. intricatus, but it is larger than 
that species and very differently coloured; the elytra at first 
appear to be uniformly coloured, but in certain lights the 
base and a postmedian space appear to be very feebly 
diluted with red. The apical half of the femora is darker 
than the basal half, on the specimen from the island being 
distinctly infuscated. The species is probably apterous. 


A specimen from the Swan River (taken by Mr. J. Clark 
from a tussock of grass) probably represents a variety of the 
species ; it differs from the type in having the head and pro- 
thorax paler (of a rather dark blood-red colour) and the elytra 
uniformly pale castaneous; the median line on the head is 
narrower (it almost vanishes in its middle), the elytral punc- 
tures are larger, and the elytral pubescence is longer and more 
upright. 


CURCULIONIDAE. 


Timareta crinita, Pasc. Numerous specimens, agreeing 
well with others from Western Australia, were obtained on 
Flinders and St. Francis Islands. On many of them the 
prothorax has denser scales, forming a fairly conspicuous vitta 
near each side; on the elytra the scales are condensed into 
numerous spots, elsewhere they thinly cover the surface and 
they are often absent from about the punctures, in conse- 
quence the elytra to the naked eye have a distinctly spotted 
appearance, although the scales are nearly always of a snowy 
whiteness (except that on the suture they are slightly darker), 
the place just beyond the incurved pogtion of the hind tibiae 
of the male is more densely clothed with long hair than else- 
where, and the middle of the incurved part appears very 
thin from some directions. 


300 


Timareta hamata, n. sp. 
Pl. xiii., fig. 4. 

3. Black, antennae and tarsi reddish. Densely clothed 
with small round greyish scales, closely adpressed to derm, 
and with numerous irregular whitish spots; with numerous 
pale, suberect setae on prothorax, and forming a regular line 
on each elytral interstice, sides and legs with longer hairs. 


Head with dense normally concealed punctures. Antennae 
long and thin, scape the length of front tibiae. Prothorax 
slightly longer than wide, sides strongly rounded, apex nar- 
rower than base, with dense normally concealed punctures. 
Elytra with shoulders strongly rounded, sides widest at about 
basal fifth, thence almost evenly narrowed to apex; with 
regular rows of large punctures, appearing much smaller 
through scales; interstices with dense and minute normally 


ee 


B D 
oe ibe Hind tibiae of He hamata, Lea, from i Le 
of view and unclothed; C D, T. incisipes, Lea; E F, 7. pilosa. 
Blackb!.°G H, 7: crinita, Pase. ; 1a ed figurata, Pasc. ; K L, 
front tibiae of T. incisipes, Lea. 


concealed punctures. Under-surface with dense punctures of 
two sizes, the larger ones scarcely concealed; abdomen with 
basal segment, widely concave in middle. Front tibiae arched 
near apex, the apex triangularly dilated on inner side; hind 
tibiae narrowed near apex, but apex itself much thickened 
and hooked, with a conspicuous fascicle of long hairs on tip 
of the hook. Length, 6-7 mm. 


Q. Differs in being wider and more convex, antennae 
shorter, seriate punctures of elytra smaller, basal segment of 
abdomen gently convex, front tibiae shorter and scarcely 
arched near apex, hind: tibiae shorter and thicker, apex itself 
wider but not hooked or fasciculate. 


Hab.—South Australia: Flinders Island (Prof. F. Wood 
Jones). Type, I. 15256. 


At first glance apparently like small specimens of T. 
pilosa, but at once distinguished by the hind tibiae of the 
males (compare figs. A B with E F). T. pustulosa has some- 
what similar ones, but the front tibiae are less swollen towards 


301 


base and the elytra are very different. Parts of the under- 
surface and of the femora and tibiae are more or less obscurely 
reddish on some specimens, but on most of them those parts 
(except as to their clothing) are black or blackish. The 
white spots are most numerous on the sides and apical slope 
of the elytra, where they are often accentuated by the 
adjacent scales being more or less sooty; on the prothorax 
the white scales usually form a distinct stripe towards each 
side, and parts of a median line, on the head and rostrum 
the scales are usually entirely white; on some specimens some 
small patches of scales are shining. 


Timareta incisipes, n. sp. 
Pl. xiii., fig. 5. 

3. Black or blackish, antennae and tarsi reddish. 
With dense, small, round scales closely adpressed to derm; 
with numerous subdepressed setae on prothorax, and forming 
a regular row on each elytral interstice. 


Head, prothorax, elytra, and under-surface as described 
in preceding species. Front tibiae trisinuate on lower surface, 
the sinus near apex appearing as a conspicuous notch; hind 
tibiae with a deep notch near apex, the notch with long hairs 
about it. Length, 5-6 mm. 

Q. Differs in being rather more robust, antennae and 
legs shorter, tibiae not notched and abdomen convex. 

Hab.—South Australia: St. Francis, Eyre, and Franklin 
Islands (Prof. F. Wood Jones). Type, I. 15257. 

The. body parts of this, the preceding species, and of 
T. crumta and T. mlosa are much alike, and the females are 
difficult to satisfactorily distinguish; but the males may be 
quickly identified by the hind tibiae alone; on the present 
species the front tibiae as well as the hind ones, are notched. 
On several specimens the under-surface, tibiae, and even 
occasionally the elytra, are obscurely reddish. The scales are 
scarcely alike on any two of the 18 specimens before me; 
they are usually of a pale slaty-brown, with more or less 
large patches, or numerous sooty spots, interspersed with 
white or bluish-white spots; on the prothorax the white scales 
form irregular lateral vittae; on an occasional specimen the 
scales are mostly sooty-brown, with numerous bluish-white 
spots; on one they are whitish obscurely mottled with pale 
brown; on two specimens many of the scales have a soft 
golden lustre; many specimens have an ochreous spot on the 
forehead. The setae on the shoulders are longer than on 
other parts of the elytra, but they are not of the great length 
of some of the sand-frequenting species. The tibiae of both 


: 302 


sexes are each tipped with a conspicuous comb-like fringe of 


setae, as they are on most species of the genus. 

Otiorhynchus cribricollis, Gyll. Black Rocks. 

Mandalotus tenuwicorns, Lea. Black Rocks. 

M. ventralis, Blackb. Flinders Island. 

Perperus languidus, Er. Flinders Island. 

Zephryne, sp. One specimen of a species evidently near 
Z. geometrica was obtained on Franklin Island; but as the 
colours of species of the genus vary considerably, it is not 
desirable to name an unique. 

Desiantha. maculata, Blackb. St. Francis Island. 

Eloeagna squamibunda, Pase. St. Francis and Franklin 
Islands. | 

Halorhynchus caecus, Woll. Two specimens of this 
curious little blind species were taken on Flinders Island ; 
it was named originally from Western Australia, but has 
been taken on Kangaroo Island and on beaches near Adelaide. 


Pentarthrocis, n gen. 


Head rather small. Eyes very small, composed of a 
few coarse facets. Rostrum moderately long, slightly in- 
curved between base and insertion of antennae, in front of 
antennae slightly wider and parallel-sided. Antennae rather 
short; funicle the length of scape, first joint slightly longer 
than second and third combined, third shortest of all; club 
indistinctly jointed. Prothorax rather elongate, sides gently 
rounded, base wider than apex. Scutellum invisible. Elytra 
elongate, with rows of large punctures in regular striae. Meta- 
sternum elongate. Abdomen with third and fourth segments 
very short, the others large. Legs rather stout; front tibiae 
with small subapical spur, and large terminal hook; tarsi 
with third joint moderately dilated, the clawjoint rather long 
and thin. 

Of the Australian genera with the funicle five-jointed 
the present genus is distinguished from Cossonideus by the 
small eyes; Halorhynchus is blind; Pentamimus and 
Pentarthrum have much shorter rostrum with much larger 
eyes; Conlonia has thinner rostrum, more parallel-sided body, 
and seriate arrangement of the elytral punctures (themselves 
much smaller) scarcely in evidence; and Microcossonus has 
much larger eyes, scutellum conspicuous, etc. In catalogues 
it should be placed near Pentarthrum. The only known 
species has somewhat fusiform outlines, and straggling hairs 
on the sides; its rostrum has a slight resemblance to that of 
some species of Cossonus. 


303 ; 


Pentarthrocis ammophilus, 0. §p. 
Pl. xiii., fig. 6. 

Dark piceous-brown, elytra sometimes dark castaneous. 
Some long straggling hairs on sides of prothorax and of 
elytra, and some shorter ones on under-surface and legs. 

Head smooth, convex, and with sparse and minute punc- 
tures. Rostrum about twice as long as its apical width; 
with rather sparse and small but distinct punctures, becoming 
more numerous about apex. Prothorax with sides evenly 
rounded and gently increasing in ‘width from apex to about 
basal fourth, and then decreasing to base; with sharply 
defined, fairly large and numerous but not crowded punctures 
on upper-surface, denser and larger on sides. LElytra at base 
wider than base of prothorax, shoulders strongly rounded, 
sides gently rounded and widest at about middle; with rows 
of large, regular punctures, in rather deep striae; interstices 
evenly convex, and each with a row of minute punctures. 
Sterna and two basal segments of abdomen with coarse punc- 
tures, smaller and more crowded ,on apical segment, and 
absent from the third and fourth. Length (excluding 
rostrum), 2°75-3°25 mm. 

Hab.—South Australia: St. Franics Island (Prof. F. 
Wood Jones); Western Australia: Geraldton (A. M. Lea). 
Type, I. 15304. 

Some specimens are almost uniformly coloured through- 
out, but on others the. elytra, club, and sometimes parts of 
the legs are slightly paler. On the male there is a wide 
shallow depression on the two basal segments of abdomen, 
on the female those segments are flat in the middle. All 
the specimens were obtained at the roots of beach-growing 
plants. 

sid COCCINELLIDAE. 

Scymnus flavifrons, Blackb. One specimen taken from 
a rat’s nest on Franklin Island. 

Rhizobius ruficollis, Blackb. Black Rocks. 


—_—_- 


' DESCRIPTION OF PLATE XIII. 
Saragus oleatus, Carter. 

Helaeus modicus, Blackb. 
Anthicus strigosus, Lea. 
Timareta hamata, Lea. 

T. incisipes, Lea. 

Pentarthrocis ammophilus, Lea. 


Fig. 


DO 99 PO 


304 


CYLINDRO-CONICAL AND CORNUTE STONES FROM THE 
DARLING RIVER AND COOPER CREEK, 


By Ropert Puuuerne, M.B., Cu.M. 
[Read September 14, 1922.| 
PuateE XIV. 


LITERATURE. 


Apart from a few records of exhibition of single speci- 
mens of these stones at scientific meetings, the first extended 
account is by :— 


1. WaLtER R. Harper, “‘A Description of Certain 
Objects of Unknown Significance formerly used by some New 
South Wales Tribes’’ (Proc. Linn. Soc. N.S. Wales, vol. 23, 
1898, pp. 420-436, pls. xii.-xviii.). 

2. R. H. Matuews, L.S., contributed a paper to Sec- 
tion F at the Brisbane meeting of the Australian Association 
for Advancement of Science, 1909, entitled: ‘‘Some Rock 
Pictures and Ceremonial Stones of the Australian A borigines’’ 
(Proc., pp. 493-498). 


3. Ropert ETHERIDGE, JUN., in the Memoirs of the 
Geological Survey of New South Wales, Ethnological Series, 
No. 2, on ‘‘The Cylindro-Conical and Cornute Stone Imple- 
ments of Western New South Wales and their Significance’ 
(pp. 1-41, pl. ix.), gives a full account of all known to that 
date on the subject, with an analysis, illustrations of many 
specimens, and a map of distribution. 

4. Eytmann, “Die Eingeborenen der Kolonie Siid- 
australien,’’ taf. xxxi., f. 1910, figures a single specimen from 
Cooper Creek with short reference. 

The early explorers of New South Wales do not mention 
these stones, and it is especially singular that neither Howitt 
nor Gason, who wrote exhaustively on the natives of the areas 
in which these objects occur, refer to them in any form. 
Howitt’s great work is so exhaustive that if anything had been 
known about the use of these stones it would certainly have 
not escaped his notice. 


Mr. Simpson Newland, who lived on the Paroo River 
from 1861 to 1876, tells me that the stones were present on 
his station, but that the natives, then very numerous, took 
no notice of them, neither using them nor avoiding them in 
any way, and had no name for them. 


305 


Mr. John Conrick, of Nappa Merrie, Cooper Creek, 
where several have been found, tells me that, although he 
has lived there since the early seventies, he has never seen 
them used or noticed by natives, and that they are known 
there simply by the name of ‘‘Moora.” Now the word 
“Moora,’’ in Gason’s Vocabulary of the Dieri of Cooper 
Creek, gives the meaning as Creator or Good Spirit, and as 
“Moora Moora’’ is frequently mentioned in legends (re- 
counted by Howitt), Sir J. G. Fraser, in his ‘‘Totemism and 
Exogamous Marriage,’’ vol. 1, points out that Gason’s mean- 
ing is erroneous, and that ‘‘Moora Moora’’ were ‘‘nothing 
more than the legendary predecessors or prototypes of the 
Dieri,’’ comparable to the Alcheringa ancestors of the Arunta 
of Central Australia. 

The significance of the foregoing seems to be that the 
objects in question are of such antiquity that their origin and 
use are lost in the past, as regards the present aborigines, 
and that any explanations they try to give are purely 
imaginary. Such explanations as these:—(1) Of use in tooth 
avulsion ceremonies (3, p. 14); (2) as a fetish to procure a 
good supply of snakes, given to Gregory (3, p. 14); (3) cere- 
monial use in connection with nardoo harvest (2, p. 497); 
(4) bora message stones (3, p. 12), show what various accounts 
aborigines will give in their desire to impart information. 
I think, therefore, that we may conclude that the aborigines 
have no knowledge, even traditional, of the origin and uses 
of the objects in question. Etheridge (3) carefully considers 
the ten suggested uses and narrows the probabilities down 
to one or two. 


GENERAL DESCRIPTION. 


At present some two to three hundred of these stones 
exist in Museum and other collections in Australia, besides 
many reported to have been sent to Germany from Menindie 
some years ago. There is no note of them in available 
German ethnological literature. z 

They are all of the same character—cylindrical from 5 to 
30 inches in length, mostly cupped at the base and composed 
of clay, kopi, sandstone, slate, or hard quartzite. Some are 
curved to form the Cornute form. The raw material from 
which they have been shaped comes from the outcrops at 
some distance from the alluvial area where they are mostly 
found, on claypans or in the blown sandhills. Sir Douglas 
Mawson says, for instance, that the slate must have come 
from as far away as Cobar or Broken Hill. Those of kopi 
are made from gypsum, with or without an admixture of 
clay, and are sometimes quite friable on the surface. The 
section is nearly always approximately circular. 


306 


MARKINGS. 

A large proportion of the stones examined present mark- 
ings, especially the softer ones. The hard quartzite specimens 
seldom, or never, exhibit them. The most common form 
of marking is what we might call ‘‘tally marks’’—small 
incisions, single, in pairs, threes, or in linear series. There 
may be as few as six, or as many as several hundreds. In 
one specimen (1, pls. xiii. and, xiv.) linear series of these 
marks have been scored through by paired, parallel, longi- 
tudinal marks, while other series are unscored. It is hardly 
‘to be doubted that these are actually tallies recording a 
number of objects or events. The keeping of tallies for 
various purposes is well known as occurring amongst Aus- 
tralian aborigines, and not unknown even amongst Europeans. 
“Broad arrow’? marks occur, and it is highly probable 
that these, as in rock carvings and paintings, indicate emu 
feet or even tracks [see illustration of rock carvings on 
Burnett River (2)|. Their use on the cylindrical stones is a 
mystery, unless we consider them the most frequent and most 
easily executed form of aboriginal decoration. 

Circular markings may occur along the length of the 
stone, or several may be present at the pointed end |[fig. 1, 
the Praeputial Rings’’ of Etheridge (3)]. These, apart from 
the hooks and stars (1, pl. xili.), certainly variants of the 
emu track marks, exhaust the forms of sculpture observed on 
the cylinders. 

Now the assigned uses of these stones are many and 
various, and have been discussed at length by Etheridge (3, 
pp. 3-18). He dismisses them all except one, or possibly 
two, as untenable. While on the slender evidence admitting 
the possibility of the snake-fetish theory, he holds the Phallic 
theory to be more tenable, in which view he is supported by 
the authority of the late Sir Edward Stirling, F.R.S., and 
Prof. J. W. Gregory. While direct evidence is unfortunately 
wanting, and@ Gason in his account of the circumcision cere- 
mony of the Dieri tribe expressly omits details, he would 
certainly have mentioned objects so striking if they had been 
in common use. It would be well if we could follow up this 
theory and see if there is any indirect evidence to support it. 
The shape of the stones is at least suggestive, and Phallicism 
is a widespread cult among primitive peoples, the world over, 
and not unknown in higher civilizations. 

Schliemann, in Ilios, figures several objects in stone and 
marble found during excavations at ancient Troy, which he 
supposes to be phalli or priapi. One of those figured on 
p. 452, No. 682, bears a striking resemblance to the one 
figured (fig. 1), even to the praeputial rings. The likeness 


307 


may only be accidental, and the marble object of the ancient 
Trojans may have been misidentified, still I mention the 
striking resemblance for what it is worth. An objection 
may be raised that the cult would have been universal in 
Australia and not confined to the central eastern area, but 
against this we have the localized Alcheringa cult with its 
equally striking stone churinga spread over a smaller area. 

If we accept the views of Churchward, now gaining the 
attention of anthropologists, that mankind originated in the 
great lake districts of Africa, we find opened up a path 
which leads to an understanding of the origin of our aborigines 
and their beliefs. In his two books, ‘‘Signs and Symbols 
of. Primordial Man’’ and ‘‘The Origin and Evolution of 
Mankind,’’ he pictures the Pygmy exodus throughout the 
world and their displacement and annihilation by the people 
of the second Nilotic exodus to which our aborigines, accord- 
ing to him, belong. He states that the Pygmies of the first 
Nilotic invasion were displaced in Australia and eventually 
only remained in Tasmania. 

The recent discovery of plateau implements in Central 
Australia by Professor Howchin (Trans. Roy. Soc. S. Austr., 
vol. xlv., 1921, p. 206, pls. xi. to xxi.), and also by Mr. 
Campbell at Millar Creek, strengthens this view, and the 
remarkable legend told in Mr. Simpson Newland’s book, 
‘“Paving the Way,’’ chap. xi., ‘‘The Doom of the Mullahs,’’ 
may be the traditional account of the fall of the Pygmies 
in Australia. At any rate, Professor Krause thought it of 
sufficient importance to give an account of the legend in the 
Zeitschrift fiir Ethnologie of the Berliner Anthropologische 
Gesellschaft, vol. 34, 1902, p. 263. 

The Pygmies who still live in Africa, New Guinea, and 
elsewhere are a non-totemic people, and seem by isolation 
to have retained their purity. This throws a new light on 
the anthropology of the extinct Tasmanians, who had the 
true peppercorn hair of the Pygmies, no totems, and no 
boomerangs. © The second Nilotic exodus brought the 
boomerang, a very ancient weapon in Egypt (vide Horus I. 
holding in left hand a boomerang, Book of the Underworld), 
also at Deirel Bahari a statue of a Prince of Punt carrying 
a boomerang (vide Churchward, ‘Origin and Evolution of 
Mankind’’), and with it the signs and symbols of the Nilotic 
people and their palaeolithic stone implements. 

Now the’ Phallic Cult originated in Egypt, where it was 
identified with the God Osiris, and from thence it was carried 
all over the world, was elaborated later by the Greeks and 
Romans, and crops up to-day in the maypole and the cere- 
monials of the Lingayat Sect, in Southern India (wde 


308 

Lingayat, ‘‘Castes and Tribes of Southern India,” vol. iv.). 
The whole account of the origin and spread of the cult is to 
be found in Rolle, Recherches sur le ‘‘Culte de Bacchus,’’ 
Paris, 1824, vol. i., p. 2. Now, in the light of this it is not 
improbable that the people of the second Nilotic exodus 
brought this rite with them, not necessarily associated with 
the ceremonial of circumcision, for in the area where the 
cylindro-conical stones are commonest circumcision was not 
practised by the aborigines in modern times. What may 
have been the condition in ancient times we shall never 
know, but I suggest that it is by following up this clue that 
our efforts of gaining further knowledge of the matter are 
most likely to be rewarded. The whole subject is bound by 
the difficulty of visualizing the enormous antiquity of man 
and his wanderings in prehistoric times. 


DESCRIPTION OF PLATE XIV. 


Fig. 1. Upper third of cylindro-conical, made of kopi, showing 
‘‘praeputial rings” of Etheridge. Nat. size. 


Fig. 2. Phallus or priapus, from Schliemann, Ilios, p. 452, No. 
682, for comparison with fig. 1. 


Fig. 3. Portion of cylindro-conical of slate, showing ‘‘tally-marks.”’ 


309 


AUSTRALIAN COLEOPTERA 
PART III. 


By Ausert H. Eston, F.E.S 
[Read September 14, 1922. |] 


HALIPLIDAE. 

I was asked to investigate the question, regarding the 
number of joints in the antennae of the Halipli, by Mr. 
Sloane, to whom I desire to express my thanks for specimens 
of exotic species, and for his kindly advice and suggestions. 

I had already prepared a drawing and notes on Haliplus 
ruficolus, De Geer (Germany), when I heard from Mr. 
Sloane that Dr. Frits van Emden had already published a 
paper (Entomologische Mitteilungen, Band xi., Nr. 2, 15 
Marz, 1922) with a drawing and a description of an antenna 
of this insect, and, as I have been able to dissect joint 1 from 
its socket in the head, I thought it desirable to publish this 
drawing in addition to the antenna of H. testudo, Clark 
(Australia). With both of the above species I was able with 
relaxed specimens to move each of the individual eleven joints 
separately, the basal joint moving quite freely in its socket 
in the head. 

In addition to those names already mentioned by Dr. 
van Emden, we find in the following publications the antennae 
of the Haliplidae referred to as having ten joints: — 
Lacordaire, vol. i1., p. 411 (Haliplus), ‘‘Antennes courtes, 
de 10 articles: 1 petit, 2-9 obconiques subégaux, 10 plus long, 
terminé en pointe.’’ Kraatz, Insecten Deutschlands, p. 9 
(Haliplini), ‘‘Antennae frontales, decemarticulatae.’’ Sharp, 
Cambridge Natural History, vol. ii1.. p. 209, “‘Antennae bare, 
_ten-jointed.”’ Packard, Guide to the Study of Insects, 
p. 436, “In Haliplus the antennae are ten-jointed.”’ Rye, 
British Beetles, p. 62 (Haliplus), ‘“‘. . . their antennae 
are ten-jointed.’”’ Sharp, in the Biologia Centrali-Americana, 
vol. i. (2), gives a figure of Haliplus solitarius (pl. i., fig. 1), 
but in the description on page 2 does not even mention the 
antennae. Stephens, Manual of British Beetles, p. 61, speak- 
ing of Haliplus, says, “antennae ten-jointed.’’ 

Apparently all these writers had regarded the two basal 
joints as one, the first division being considered the “‘bulb 
of insertion,’’ similar to that found in the Carabidae. The 
insects comprising the genus Haliplus have no bulb to the 
first joint (fig. 1, a and c), which is inserted into the head 
and moves freely in its socket (fig. 1, b), and joint 2, in turn, 
articulates on joint 1. For the purpose of comparison a 


310 


drawing is given of an antenna of a carab, Lebimorpha 
benefica, Newm. (fig. 1, d), showing the bulbous basal part 


Kies tf. 
a, Antenna Haliplus ruficollis, De Geer; b, socket for 
reception of antenna H. ruficollis; c, antenna H. 
testudo, Clark ; d, antenna Lebimorpha benefica, Newm. 


of joint 1; on the first joint of each antenna is to be seen a 
long tactile seta situated in the middle before the apex. 


PAUSSIDAE. 
ARTHROPTERUS ARTICULARIS, Elston. 


The length of this species should read 9-95 mm., not 
5-5°5 mm., as printed. 


HISTERIDAE. 
CHLAMYDOPSIS EPIPLEURALIS, Lea. 


Five specimens of this species were taken by R. F. Kemp 
and myself from the nest of the common small black ant 
(Iridomyrmex, sp.), in the Mount Lofty ranges. They are 
variable in size, ranging from 2°5 to 4 mm. in length; the 
smallest is much paler than the typical form, its colour is 
testaceous, with parts of the elytra almost flavous. | 


COLYDIIDAE. 


Todima fulvicincta, n. sp. (Fig. 2). 

Elongate; piceous, with clypeus, antennae, sides of pro- 
thorax, portions of elytra, and parts of legs, fulvous. 
Scantily clothed with short, golden hairs, fairly numerous on 
front of prothorax, and on the elytra arranged in rows towards 
apex. Under-surface nitid, piceous, except forepart of head 
and sides of prosternum, which are fulvous; sparsely clothed 
with short, depressed, golden hairs. 


(1) Elston, Trans. Roy. Soc. S. Austr., 1919, p. 342. 


311. 


Head subquadrate, anterior margin and sides near the 
middle contracted, with a shallow, elongate depression near 
base of each antenna; and with dense, small, subrugose punc- 
tures. Antennae about four-fifths the length of head, moder- 
ately robust, second joint approximately twice the length of 
the first, joints 4 to 8 little more than half the width of the 
second, and not quite as long, the ninth wider than the 
eighth, the tenth more than twice as wide as the ninth and 
almost semicircular in shape, the apical longer than and 
_ about three-quarters the width of the tenth, almost circular. 


Todima fulvicincta, n. sp. A, front leg. B, antenna. 


Prothorax about one and half times wider than head, the 
anterior margin wider than the base, sides contracted near 
the middle, the anterior angles acute, posterior ones rounded, 
disk with a large, shallow, obovate depression, and divided. 
transversely with a more or less distinct raised portion; with 
dense, subrugose punctures, larger and more distinct than 
those on head. Scutellwm very small and somewhat semi- 
circular. Hlytra at base slightly wider than prothorax, and 
about three times as long, sides parallel to beyond the middle, 
and evenly rounded towards apex; with closely placed seriate - 
punctures, larger than those on prothorax. Legs robust, first 
two joints of tarsi dilated. Length, 3°5-4°5 mm. 
Hab.—South Australia, taken in Xanthorrhoea on the 
summit of the Devil Peak, near Quorn (R. F. Kemp and 
A. H. Elston). Type, in author’s collection; co-type, I. 
15232, in South Australian Museum. 

A very distinct species, and easily distinguished by its 
markings. The fulvous part on the prothorax is widest in 
front, sometimes disappearing before base, and on each elytron 
is in the form of a crescent, the convex side reaching a little 
more than half-way across, between the margin and the 
suture; this crescent-shaped part varies somewhat in size on 


312 


the twenty-two examples before me; a narrow edge at the 
apex of the elytra is also fulvous, and on most specimens is 
joined to the crescent-shaped patch with a very narrow strip 
at the margins. The head and prothorax are in parts 
shagreened owing to the density of the punctures. The femora 
and tibiae are brown, in parts paler, the base and apex of the 
latter, and the tarsi are fulvous.. A more robust species than 
T. lateralis, Blackb., with the shape of the prothorax very 
different, the punctures on the elytra larger, and the two 
first joints of the tarsi more dilated. 


CLERIDAE. 
Phlogistus agraphus, 2. sp. 

Upper-surface piceous, subnitid, apendages of mouth and 
the antennae testaceous, club of latter infuscated, head in 
parts reflecting blue, legs dark blue to piceous. Clothed with 
moderately long griseous hairs, thicker at the sides of pro- 
thorax than elsewhere. Under-surface green, with brassy 
reflections, and scantily clothed with griseous hairs. 

Head with a distinct, round, moderately deep fovea 
between the eyes, and with closely-set, somewhat deep punc- 
tures, more or less rugose towards forepart. Antennae reach- 
ing to about middle of prothorax, joints 9 and 10 transverse, 
the eleventh ovate-acuminate. Prothorax transverse, the 
anterior margin wider than the basal one, before apex with 


a curved, and at the base with a straight transverse Im- ~ 


pression, the centre of disc with a moderately deep depression, 
in the centre of which is a tolerably long, deep furrow, the 
sides are strongly rounded, the greatest distance between 
them near the middle; with somewhat dense punctures, about 
same size as those on head but more feeble, transversely 
rugose on disk and sides. Hlytra at base distinctly wider than 
prothorax, and about twice as long as wide, sides parallel 
to beyond the middle, then gently rounded off towards apex ; 
with ten rows of large, deep, quadratic punctures, which start 
from behind the base and extend to about the apical quarter 
of elytra, the apical fourth with rows of almost obsolete 
punctures. Legs robust, posterior femora almost reaching 
apex of abdomen. Length, 9°5-11 mm. 

Hab.—Western Australia: Cottesloe (H. M. Giles); 
Perth (J. Clark). Type, in author’s collection; co-type, 
I. 15337, in South Australian Museum. 

A very robust species; on some specimens the greenish 
reflections on the elytra are stronger than on others; on the 
elytra the basal and apical portions are more nitid than the 
remainder, the large seriate punctures suddenly cease at the 
apical fourth, then continued, only very feebly, in rows to 
the apex. In sculpture it comes nearest to Ph. imperialis, 


May 


5 

a 
} 

NY 


313 


Gorham, but differs in being more robust, in the shape of 


the prothorax, the punctures on same more feeble, the basal 
part of elytra more tumid, and the punctures on elytra some- 
what larger. ; 
_ Phlogistus rubriventris, 1.sp. 
Shining black, palpi, apical joint of antennae and tarsi 
slightly diluted with red, the abdomen and tarsal claws red ; 
moderately clothed with pale hairs, semi-erect on the upper- 


surface and depressed underneath. 


Head somewhat elongate; with a large, round, interocular 
depression, and dense punctures, which are individually dis- 
tinct on the top of head, but smaller and more rugose towards 
the forepart. Antennae reaching to about the middle of 
prothorax; club three-jointed, ninth joint obconical, tenth 
almost transverse, and the eleventh ovate-acuminate. Pro- 
thorax transverse, before the anterior margin with a curved, 
and at the posterior one with a straight transverse impression, 
a moderately deep fovea on the disk, situated immediately 
behind the anterior transverse impression, and a shallow 
depression at each side near the middle; the lateral margins 
are well rounded, the greatest width between them being 
near the middle; less closely punctured than the head, the 
punctures are somewhat scattered on the disk, but at the 
sides they are closer and more or less rugose. Hlytra at base 
much wider than the prothorax, about twice as long as wide, 
sides almost parallel and gently rounded off towards apex, 
shoulders prominent; with ten rows of moderately large, 
almost quadratic, punctures, which begin at the base and 
extend to the extreme apex. Legs comparatively short, the 
posterior femora not reaching the apex of elytra, claws moder- 


ately long, with a conspicuous tooth, situated on the inside 


near the middle. Length, 7-8°5 mm. 

Hab.—Western Australia: Eradu (J. Clark). Type, in 
author’s collection; co-type, I. 15338, in South Australian 
Museum. 

This species is very distinct from any other Phlogistus 
known to me, the very conspicuous median teeth on the claws 
made me, at first, feel doubtful as to it being a Phlogistus, 
but on examining the claws under a moderately high power, 
I find that these teeth appear to have their origin at the base 
of the claws. The punctures at the base, on the shoulders, 
and towards the apex of elytra are slightly smaller than those 
on the disk, but nevertheless, are very distinct, the extreme 
apex of elytra is truncate, and at the sutural angle somewhat 
acuminate. A specimen from New South Wales, in the col- 
lection of Dr.-E. W. Ferguson, is possibly a variety of this 
species; it differs from the type in having the palpi and 
antennae pale, the apical joint of the latter more elongate; 

3 | 


314 


the prothorax is somewhat differently shaped, in rwbriventris 
the anterior and posterior margins are about equal in length, 
but in the New South Wales specimen the anterior margin 
appears to be wider, also the surface of the prothorax is less 
nitid, not so uneven, and with more feeble punctures; other- 
wise it agrees very well with the above description. 


Phlogistus ungulatus, 0.Sp. 


Black, subnitid, antennae and appendages of mouth | 


brownish, claws reddish. Somewhat thickly clothed with pale 


hairs, more or less shaggy on the upper-surface and depressed — 


on the under-surface. 

Head with a shallow longitudinal impression near the 
base of each antenna, and with small, shallow punctures, 
somewhat scattered on the top, but towards forepart closer, 
and more or less rugose. Antennae short, barely reaching 
to middle of prothorax, club three-jointed, joints 9 and 10 
transverse, the apical almost as long as the two preceding 
combined and obtusely pointed. Prothorax barely transverse, 
behind the anterior margin and at the base with compara- 
tively shallow transverse impressions; a feeble longitudinal 
impression on the disk, situated immediately behind the 
anterior transverse one, and on each side near the middle 
of the lateral margin a round depression ; the punctures are 
somewhat more feeble than those on the head, and rugose 
at the sides. Llytra at the base distinctly wider than pro- 
thorax, and about twice as long as wide, sides parallel to 
beyond the middle and then gently rounded off towards apex ; 
with ten rows of moderately deep and almost quadratic punc- 
tures, starting at the base and reaching to the extreme apex. 
Posterior femora do not reach apex of elytra, the basal teeth 
on the claws very long and conspicuous. Length, 4°5-5°5 mm. 

Hab.—Western Australia: Swan River (J. Clark). Type, 
in author’s collection. 

Very closely related to the preceding species but easily 
distinguished from it by its smaller size, the whole of the 
under-surface is black, more hairy, and the punctures, par- 
ticularly on the prothorax, are more feeble, and with apex 
of each elytron rounded. The peculiar structure of the claws 
readily distinguishes this, and the preceding species, from all 
previously described ones, the basal teeth on the claws of 
the present species are very elongated, nearly extending to 


the apex of the claw, and giving it the appearance of being ~ 


cleft. 
Phlogistus leucocosmus, D.Sp. 
Upper-surface subnitid, blue, antennae and apendages 
of mouth more or less testaceous, head greenish-blue, elytra 
almost violet, clothed with somewhat shaggy pale hairs, very 


q 


315 


densely arranged near middle of elytra, and forming an 
oblique fascia on each. Under-suriace greenish- -blue and 
rather scantily clothed with pale, depressed hairs. 

Head wide, with a large round interocular depression 
and close rugose punctures. Antennae moderately long, reach- 
ing to beyond the middle of prothorax, joints 9 and 10 
obconical, the eleventh ovate-acuminate. Prothorax almost as 
long as wide, before the apex with a curved, and at the base 
with a straight transverse impression, the latter deeper than 
the former, the disk with a deep round depression, the top 
of which touches the anterior transverse impression, sides 
well rounded, the greatest distance between them being near 
the middle ; middle of disk with fine transverse wrinkles, the 
punctures only individually distinct near apex and sides. 
Elytra at base wider than prothorax and about twice as long 
as wide, sides almost parallel to beyond the middle then 
gently rounded off towards apex, humeral angles prominent, 
with ten rows of moderately large punctures, which begin 
from behind the base and end abruptly at the median fascia 
of hairs, the base with only a few small, scattered punctures, 
the posterior part behind the fascia with disjointed rows of 
obsolete punctures. Posterior femora do not reach apex of 
posterior part of body. Length, 6°5-7 mm. 
| Hab.—Western Australia: Swan River (J. aoe 
Type, in author’s collection. 

A very distinct species, and readily distinguished f the 
oblique fascia of pale hairs near the middle of the elytra. 
On one specimen the head is green with brassy reflections, 
and underneath the fasica of hairs there are traces of green. 
The sculpture of the elytra is very similar to that of Ph. 
mundus, Blackb., but is distinguished from it by its colour 
and the elytral fascia, the shape and puncturation of the 
prothorax is also different, and the eyes are somewhat more 
‘prominent. 

PHLOGISTUS PUNCTATUS, Hintz. 

A specimen from Bowen, Queensland, agrees very well 
with the author’s description, except that the whole of the 
antennae are testaceous, the labrum, anterior and inter- 
mediate legs are also of the same colour, the two latter have 
their knees infuscated, the posterior tibiae on the under- 
surface are pale. The sutural row of punctures begins almost 
immediately behind the scutellum. 


TARSOSTENUS UNIVITTATUS, Rossi. 
Omlo incertus, Macl. 
Macleay’s name will now have to be added to the several 
- synonyms of this cosmopolitan species. There are specimens 
of it in my collection from Queensland, South Australia, 
ae Australia, and they are, inter se, variable both 
| 


316 


in size and colour. A specimen from South Australia is 
much paler than the typical form, its colour is a reddish- 
brown with the head almost black, and the fascia on the 
elytra yellow; on some the whole of the legs are ferruginous, 
here and there infuscated. 


Tarsostenodes leucogramma, 0.Sp. 


Elongate; testaceous, with a spot on each elytron near 
_ the scutellum, a larger one below each of these, the posterior | 
half of elytra, and parts of the legs, bluish-black or black; 
a little behind the middle of elytra are two raised white bands 
obliquely placed, touching the margins but not the suture, 
midway between these and the humeral angles, near but not 
touching the margins, two raised white maculae, and about 
midway between the latter, near the base but not touching © 
the suture, two similar, but somewhat smaller, maculae. 
Clothed with moderately long, semi-erect, black hairs, those 
on the posterior part of elytra are thickly interspersed with 
shorter and more depressed silvery ones. Under-surface 
testaceous, with the exception of the abdomen, which is 
black; very scantily clothed with short pale hairs. 

Head with small, closely placed, rugose punctures. 
Antennae slender, second joint small and_ globular, 
3 to 8 elongate, the eighth distinctly shorter than the pre- 
ceding one, club three-jointed, apical joint ovate-acuminate. 
Prothorax elongate, convex, with a shallow subapical trans- 
verse impression, posterior margin narrower than the anterior 
one, sides rounded near the middle; with closely placed punc- 
tures, somewhat larger than those on head and more indi- 
vidually distinct. Scwtellum small and subtriangular. Hlytra 
distinctly -wider than prothorax, about three times as long 
as their width at base, sides parallel to about the middle, 
then slightly dilated, with rows of moderately large, reticulate 
punctures, beginning at the base and ceasing abruptly at the 
post-median white fascia, apical portion with very small, 
almost obsolete punctures. Legs long and somewhat slender. 
Length, 4°5-5°5 mm. . 

Hab.—Queensland: National Park (H. Hacker); New 
South Wales: Illawarra (W. du Boulay). Type, in author’s 
collection; co-type, I. 15336, in South Australian Museum ; 
and co-types in Queensland Museum. 

Apparently a variable species in its colour and markings, 
for on some the prothorax is much darker, the lateral mar- 
gins and base being almost black; two specimens have the 
anterior part of the elytra entirely pale, with the four white 
maculae more or less distinct; the humeral angles are either - 
black or testaceous, and the black portions of the anterior 
part of elytra are sometimes at the margins joined to the 


317 


black posterior part, and the latter, on account of the faint 
sculpture, is more nitid than the remainder of the elytral 
surface. 

Eleale aenea, 10.Sp. 

Whole of upper-surface coppery, nitid, three apical joints 
of antennae dull black, labrum with greenish reflections. 
Clothed with long, black, erect hairs, more numerous on sides 
and legs, where they are interspersed with white ones, 
scutellum very scantily clothed with white pubescence. Under- 
surface blue with greenish reflections, intermediate and pos- 
terior coxae violet, clothed with long, shaggy, white hairs, 
thicker at the sides than elsewhere. 

Head well produced in front, with a deep almost circular 
depression between the eyes, and a more elongate one at the 
base of each antenna; with close, fine punctures, individually 
distinct on top and confluent towards forepart. Antennae 
reaching to the base of prothorax, club five-jointed, joints 9 
to 11 compressed, the apical one on the inside with a large, 
but not deep, emargination, the outside rounded and with 
the apex acute. Prothorax slightly narrower than the head 
(including the eyes), longer than wide, subapical transverse 
impression almost obsolete, subbasal one more distinct, sides 
near the middle evenly rounded; disk flat, with a shallow 
depression in the middle immediately in front of the base, 
and one on each side near the middle; near apex with fine, 
more or less distinct punctures, elsewhere transversely wrinkled. 
Scutellum almost circular and minutely punctured. Llytra 
at base wider. than prothorax and about thrice as long, sides 
straight and parallel nearly to apex and then evenly rounded, 
humeral angles slightly salient, and behind scutellum with a 
large, round, and comparatively deep depression ; with close, 
moderately large, deep, reticulate punctures, here and there. 
transversely confluent, smaller at base and apex, but never- 
theless quite distinct. Legs somewhat slender, posterior 
femora not reaching apex of elytra. Length, 8 mm. 

Hab.—South Australia: Myponga (R. F. Kemp and A. 
H. Elston). Type, in author’s collection; co-type, I. 15248, 
in South Australian Museum. 

Distinguished from £. aspera, Newm., by having the 
sides of the prothorax dilated near the middle, the transverse 
wrinkles on same coarser, and the punctures on elytra less 
crowded and more reticulate. Very near FH. reicher, Spin., 
but with the antennae more slender, transverse wrinkles on 
prothorax somewhat finer, and punctures on elytra much 
smaller and more crowded. In scultpure very similar to 
H. wridis, Guerin, but distinguished from it by its colour, 
the club of the antennae more distinctly five-jointed, and. the 
transverse wrinkles on prothorax finer. 


318 


ELEALE SIMPLEX, Newm. 


Specimens from Western Australia differ from the typical 
form in being larger, more greenish in colour, somewhat less 
nitid, and in having the antennae dark with the first three 
or four joints more or less testaceous; on one, an inter- 
mediate form, joints 1 to 4 are testaceous, 5 to 8 are dark, 
here and there paler, and the three apical joints are a sordid 
testaceous. leale wntricata, Klug, I believe to be only a 
variety of this species. 

Hab.—South Australia, Victoria, Tasmania, Western 
Australia. 

ELEALE PULCHRA, Newm. 

Two specimens from Cottesloe, Western Australia, have 
the whole of the antennae dull black, with only joints 2 and 3 
slightly tinged with red; on one the prothorax has a distinct, 
interrupted, longitudinal median carina, on the other it is 
much less distinct. This is, apparently, the form Spinola 
named #. bimaculata. - 


LEMIDIA ALTERNATA, Lea. 

Four specimens from Queensland differ from the typical 
form by the size and shape of the elytral markings. The 
‘red basal band is narrow, the submedian black band very 
wide, the postmedian red one about half the width of the 
preceding dark one, and the apical black portion about two- 
thirds the width of the preceding red part. On all of the four 
specimens the submedian black band is by far the widest. 
The whole of the legs are pale, except the posterior tarsi, 
which are more or less infuscated. 


ALLELIDEA BREVIPENNIS, Pascoe. 

A specimen taken near Ballarat, Victoria (near type 
locality), differs from the author’s description by having all 
the tarsi blackish. Pascoe in his Latin description says, 
‘‘tibiis flavis,’? and in his English delineation says, ‘“‘tarsi 
yellow.”” This may, perhaps, be an error, ‘‘tibiae yellow’ 
being meant, but only a reference to the type, which is in the 
British Museum, will definitely reveal this. The specimen 
before me has all the tibiae flavous, and the tarsi blackish. 


CURCULIONIDAE. 
MANDALOTUS LUTOSUS, Lea. 

Four specimens of the above species were taken by R. F. 
Kemp and myself from moss on the summit of Mount Lofty, 
South Australia. The male differs from the author’s descrip- 
tion in having the carina on rostrum distinct, the granules 
on the prothorax transversely arranged, the under-surface 
of body diluted with red, particularly the last two segments 
of the abdomen, the coxae and parts of the under-surface 
of legs red. 


319 


RESEARCHES ON THE INSECT METAMORPHOSIS. 


PART !1.-ON THE STRUCTURE AND POST-EMBRYONIC DE- 
VELOPMENT-OF A CHALCID WASP, NASONIA. 


PART i!.-ON THE PHYSIOLOGY AND INTERPRETATION OF 
THE INSECT METAMORPHOSIS. 


By OW. Vines, MSc... . 
Zoology Department, University of Adelaide. 


[Read October 19, 1922. | 


PLATES XV. To XXX. 


Page 
INTRODUCTION ... as ba ds ra i, hes ven, (Oe 
; Parr, 1. 

ON THE STRUCTURE AND Post-EMBRYCNIC DEVELOPMENT OF A 
Cuatcip Wasp, Nasoma ... ye vp ota is | 26 
A. The External Features _... ic ts aye pee 8) 
General Remarks ... : ‘ee cag spe, S20 
The Head and its sees: re ee i. 29 
The Constitution of the Head . BT. Ce aanh tee 
The (true) Thorax and its Decade Las eae 37 
The Abdomen and its pee id ee vie ott 
The Ovipositor sat 2 du x i eAo 
The Penis ae ot eto 
B. The Integument fee siclowieal Tewalonmerty 4 348 
The General Body Integument be ze ... 348 
Destruction of the Body Integument ; Np oan 
Metamorphosis of the Underlying Samnaidenne vag OO 
Renovation of the Epidermis ... Pe? be ome 3), 
Formation of Body Sculpturings ... aye cae Ok 
Formation of Bristles As we re es 39” 
Formation of Pubescences ... ou uk ene tes 
The Phragmas sie ile ai As Hem, SOS 
The Legs an ae aoe ae 3.5: sa) BoA 
The Wings _... “ Da ie a) aoe 
The Mouth Meo cndanes a ie me is) OO 
The Ovipositor #44 slat ibe ate Seals aoe 
The Penis ne 3 is ae ep Me! Od 
The Antennae ae Hoe so A Cae ie. Tt! 
The Organs of Vision... Cape Mes fers ee eh SOO, 
The Compound Eyes i se Sul Ba) ED 


The Ocelli Ave ny fae pis fe: Reyes oe! 


320 
Page 
C. The Respiratory System ... ue sue ... 386 
The Larval Respiratory by aieeh A. J. ... 386 
The Destruction of the Larval Tracheae ...  ... 391 
The Regeneration of the Tracheal sy Rae ... ode 
D. The Muscular System ‘igh ... aoe 
The Anatomy of the Larval Museu Sy tera .. soon 
The Structure and Post-embryonic Development of 
the Larval Muscles v4 ag hai ... 400 
The General Body Npecalahure. mp a ... 400 
The Dilators of the Pharynx aps He ...* 403 
The Destruction of the Larval Musculature ... 404 
The Dilators of the Pharynx Oe ache ... 404 
Thoracic Muscles et at Ai = wo) AOS 
Muscles of the Abdomen ... oo. 404 
The Longitudinal Abdominal Mages . Ae 
The Vertical Abdominal Muscles ie «406 
The Regeneration of the Muscular System ... ..- 408 
The Superficial Longitudinal Abdominal Musee? 409 
The Vertical Abdominal Muscles ... Es. fo. ED 
The Dilators of the Pharynx oe won URE ee 
Muscles of Mouth Appendages .... Bi tc, sae 
The Leg Muscles Me as wale as ... Al6 
The Muscles of Ovipositor ... 24 oe ee 8 
The Muscles of Flight #y. ry mes me ls: 
Intestinal Muscles ... ee ee ot ass 4D 
The Muscle Insertions 2B Bi, ee ~ony 
The Structure of the Adult Muscles ... fs sca ABG 
K. The Intestine and Related Structures ... 428 
The Anatomy and Structure of the Twtdetiaes of ibe 
Adult a : ne ie ae | ae 
General Mor pholoue p.. a be ... 428 
Histological Sirueenee: ae fie ae re 
Oesophagus Me ray wih ne ... 429 
The Crop ... a A an an ... 430 
The Gizzard A hs ie Pa vom 00 
The Stomach sas a ‘a tog op ee 
_ The Small Intestine oe, a lg jen) ae 
The Rectum a = Ag +7 ... 4381 
The Salivary Gland ... 0 J). 0.) 
The Malpighian Tubes ... me woe | eS 
The Intestine of the First Larval Tnedar ay aes a 
General Morphology ... a i) ies ... 433 
Histological Features Aah oe xt woe, 40D 
The Buccal Cavity aD Be oe ... 435 


The Oesophagus ... 03 te ie .. 435 


(oy) 
i?) 
feat 


Page 

The Midgut nee BS es at ... 436 

The Rectum ‘ we aa a ... 436 

The Salivary Glands ea nae bys ... 4386 

The Hepatic Caeca ome 4) fot 

The Post-embryonic Development of ‘tee Tate 437 
Metamorphosis of Foregut . 437 


Metamorphosis of the Widens acd aie ies hie 
ment of the Post-oesophageal part of the 


Foregut bs oa ts Nee 
Metamorphosis of Bis ia aot se us 1,1, 444 
The Rectal Glands ee ie ” ante 
The Malpighian Tubes ... wee ae na AAC 
The Salivary Glands ... ete me oe aAg 
General Remarks ... bie mi ti, aa) 
‘F. The Ductless Glands ee ve ie a i jayiAod 
The Oenocytes.... co ee Bebe) = cs Soe be Aaue 
The Lateral Intestinal Glaiaids #4 ee ogi wn, 404 
The Dorsal Abdominal Glands a es an Wee 1595) 
'G. The Fat-body es ms Aas 
Structure and Me micah locie a Fatcbody ba 4. 456 
Function of the Fat-body si hie £4 i 1 AOL 
fo The) Gonadsivy.. 2! rs ey xe! a ey a ABD 
Male Organs ast ae doe wep me eh eAGD 
Female Organs... ny wo a6 nig aD 
I. The Nervous System sede be say ny ean ATD 
Introduction inp A472 

The Ventral Nerve Cord tie Horetal Mates of 
the Nasonia Larva ... nen ae “ht i £3 

The Post-embryonic Development and Meia 
morphosis of the Ventral Nerve Cord ... spa age 2 
The Brain: its Structure and Metamorphosis... 478 
J. The Vascular System ... Aa: a x et een eee 
The Blood ... oe ae go st ee i) SAS4 
The ‘‘Heart’’ ae: ug! a, any LAY 
Structure of the ae Beart id £% gt ASS 
Metamorphosis of the Heart a: Es ... 489 

K. Appendix— 

The Degeneration Processes of the Larval Cells of 

Neasonma 7h ae cs bi sag as 8 2490 
Parr II. 

On THE PuHysIoLOGyY AND INTERPRETATION OF THE INSECT 
METAMORPHOSIS Wn aes at re , ... 492 
Summary ce ae es in BS “ash ... 504 


Bibliography ee ad; ou ee ma iat ees ha E 


322 


INTRODUCTION. 


The insect transformation presents one of the most in- 
teresting of the many phenomena of living things about us. 
To the popular imagination it is a manifestation of the super- 
natural. To the biologist it offers unrivalled material for 
the study of several fundamental tissue reactions: extensive 
tissue degenerations followed by correspondingly great tissue 
regenerations; delayed cell differentiation and cell regenera- 
tion, and sometimes even, it seems, cellular dedifferentiation ; 
while the cases of phagocytosis at times met with are extra- 
ordinary. Nevertheless, its study has been very neglected. 

Numbers of the great early anatomists—Malpighi, 
Swammerdam, Lyonet, Diirckheim—turned their attention 
to the structure of insects, and though they were able to show 
that the larvae of insects had already the same general 
anatomy as had the adult insects, yet the difficulties of the 
dissection of the soft semi-fluid contents of the pupal shell 
proved so great, that the process of transformation was not 
elucidated. 

Réaumur, it is true, had been able to show that the limbs 
of the adult insect were to be found invaginated beneath the 
surface of the body of the nymph. Newport (1832) had 
observed the concentration of the ganglia of the ventral 
nerve cord as it changed from the larval to the imaginal 
condition, but beyond these facts nothing was known; and 
Oken, who wrote his voluminous ‘‘Allgemeine Natur- 
geschichte” at about this time (1836), summarized his know- 
ledge of the process thus (vol. 5, p. 714):—‘‘At the last 
moult the insects become covered by a horny shell, which is 
devoid of feet and oral appendages. Consequently in this 
stage they lie quiet for several weeks, often throughout the 
whole winter, without feeding or moving, and in this con- 
dition are spoken of as pupae or nymphs. Under this shell 
is gradually formed the perfect insect, the fly with its three 
body parts, with its new feeding organs, feet and wings; 
finally the skin splits dorsally, the insect creeps out, waits a 
few minutes till it has hardened, and then crawls or flies away, 
to seek other food or to reproduce. This gradual step-like 
development is spoken of as a transformation or meta- 
morphosis.”’ ; | 

It was not till the publication in 1864 of Weismann’s 
great memoir on the metamorphosis of the blow-fly that any 
light was thrown on the process. Weismann, without any 
modern technique available to him, and using only the old 
method of hand dissections, studied the process with remark- 
able accuracy. His observations were made more on broad, 
general, anatomical lines. He was able to show that the 


ald 
we 
‘- 


323 


larval tissues underwent a process of disintegration— 
“‘histolysis’’ he called it—into rounded bodies, which he called 
Koérnchenkugeln, and that the imago in turn was formed 
from small areas of cells, which Swammerdam had already 
discovered, though he had not recognized their significance ; 
to these he gave the name ‘‘imaginal discs.’’ He was able 
to demonstrate the sexual organs in a young condition in 
the larva, and to show that the insect metamorphosis was 
entirely different from the alternation of generations that 
occurred in some groups of animals and plants. He demon- 
strated the occurrence of metamorphosis in most of the organs 
of the body, including the heart and nervous system, which 
other investigators with more elaborate technique at their 
disposal have since questioned; and though his observations 
were necessarily incomplete, and did not extend largely to 
cell changes, yet his conclusions were, in the main, correct. 


Since Weismann’s memoir the blow-fly (Calliphora) has 
been used by a number of investigators for the study of 
metamorphosis, so that our knowledge of the process in this 
insect, though still very incomplete, is much fuller than that 
of any other. In 1876 the. Russian Ganin wrote upon it, 
and described the imaginal “‘nests’’ within the intestine. 
In 1884 Van Rees, and in the following year, quite inde- 
pendently, Kowalevsky, guided by Metchnikoff’s recent dis- 
covery of the phagocytic action of leucocytes, showed that the 
larval tissues were destroyed by the interference of these 
colourless corpuscles of the blood. A special interpretation 
was therefore placed on Weismann’s histolysis, and the 
‘““Kornchenkugeln’’ proved to be nothing but gorged phago- 
cytes, a fact the truth of which Metchnikoff had himself 
already perceived from the drawings given by Ganin. 


Since that time a number of other observers have added 
details to the knowledge accumulated by the earlier workers: 
—Van Rees studied it in 1888; Lowne’ published a few 
observations (mostly incorrect) in 1890-1895; Vaney wrote 
about it in 1902; while Pérez published his very detailed 
work in 1910. 


In 1899, and later in 1901, Berlese published his observa- 
tions, and seriously questioned the important réle which the 
leucocytes were believed to play in the removal of the larval 
tissues. From the earlier writings it seemed to follow that 
the leucocytes attack the living tissues, so that metamorphosis 
is, in part, brought about by more than usually highly- 
endowed leucocytes. Berlese denied this conception entirely. 
As he appears to have been misunderstood by others, it is 
best to quote his own words (1901) :—‘‘Phagocytosis never 


_/ \ wrong; phagocytes play a large part in the removal of larval 


324 


occurs, and amoebocytes only become active when the muscle 
has disintegrated through internal causes.’’ By phagocytosis 
he evidently means the phagocytosis of living tissues; and 
his ‘‘amoebocytes’’ appear to be a congregation of various 
kinds of embryonic cells and leucocytes, though he does not 
specially mention these. Pérez (1910), on the other hand, 
has taken precisely the opposite view, and regards the 
leucocytes as playing the main part in the destruction of © 
tissues. ‘‘I think I have proved satisfactorily that the dis- 
integration of the muscle is due to phagocytes, and that there 
is no spontaneous fragmentation of this organ into sacrolytes, 
as Berlese thought.’’ I may say at once, that the study of 
the metamorphosis of NVasoma has led me to conclude that 
while neither statement is quite correct neither is wholly 


tissues, but such tissues are always dead. 


Besides the observations of these workers, others have 
been made on portions of the metamorphosis of ‘other insects, 
but nothing so extensive as those made on the blow-fly exists. 
In 1875-1878 Kiinckel d’Herculais published his studies on 
the structure and transformation of the syrphid fly V olucella; 
Deegener in more recent years has studied the transformation 
of the intestine in a number of insects; and Verson (1898) 
examined it in the silkworm. Pérez (1902) examined por- 
tions of the metamorphosis of the ant Formica rufa; Bauer 
studied the transformation of the brain in several insects; 
and in 1912 Giinther investigated the development of the eye 
in Dytiscus. In 1910 Poyarkoff published his very interest- 
ing observations on the metamorphosis of a beetle, Galeruca; 
he showed that, while some organs underwent the usual type 
of phagocytic histolysis, others (the integument and part of 
the intestine) passed through a remarkable process of cellular 
rejuvenation. 

It may be said then, that while we possess a considerable 
knowledge of the main features of insect metamorphosis, on 
some of the fundamental facts much difference of opinion 
prevails. Why do the larval tissues disappear? Do the 
phagocytes kill them, or do they merely remove them after 
they have died? If the latter, then how is their death brought 
about? If in one insect phagocytic histolysis occurs, and in 
another merely cellular rejuvenation, how are we to correlate 
the processes? It is these questions that I shall attempt to 
answer in the present paper. The histological changes under- 
gone by some of the larval organs, moreover, have never been 
examined—heart, peripheral nerves, ventral nerve cord, and 
others ; whilst the greatest differences of opinion prevail about 
the details of other organs such as the muscles and intestine. 


325 


An equally interesting question is the relation in which 
the insects which show a metamorphosis stand to those in 
which it is absent; this question has been discussed by Lub- 
bock (1874), and more recently by Deegener (1909). Lub- 
bock’s conclusion, that the metamorphosis was made necessary 
by the larvae developing different feeding habits and con- 
sequently different mouth-parts from those of the adult insects, 
is not very satisfactory. While it is true that the transition 
from one to the other would have to be slow and would have 
to take place during a resting stage, it fails to account for 
the metamorphosis of structures of almost negligible import- 
ance, such, for example, as the fine somatopleural membrane 
beneath the integument. It fails also to explain the meta- 
morphosis of the feeding organs in insects in which the larvae 
and adults have the same feeding habits, such, for example, 
as many of the carnivorous and leaf-eating beetles. More- 
over, the real thing to show is why the larval form should 
ever have been evolved, necessitating the parallel evolution 
of a metamorphosis, when some insects, very successful in 
the struggle for existence, have got on so well without it. 
The conclusion of Deegener, that the larval form is a stage 
graduallyinserted between the earlyembryo state and the adult, 
is undoubtedly quite correct, and seems to be usually accepted 
to-day. Nevertheless he throws no light on the reason why 
such a form should ever have been evolved, nor does he 
explain why it later transforms itself into the mature insect. 

It was to answer these several questions that the present 
work was undertaken. The insect which I have employed is 
a small chalcid wasp, Vasonia brevicorms, very common in 
Australia and America as a parasite on exposed pupae of 
muscid flies. According to Mr. A. A. Girault it is identical 
with NVasona abnormis, Boheman, from Europe, and is evi- 
dently of world-wide distribution. As the work proceeded 
I found myself at a disadvantage in that very little was 
known about the internal anatomy of chalcid wasps, while 
the study of the anatomy of the larvae had also been 
greatly neglected, and more than one very serious 
misinterpretation have been accepted as fact. I have 
therefore resolved to extend the scope of the paper. In 
the first portion the various organs of the larva 
and adult are described and a fairly detailed account of 
them is given as they transform from the larval to the adult 
conditions. In the second part I shall attempt to explain 
the physiological basis of the metamorphosis, and to discuss 
the factors which have underlain the evolution of the process. 

The earlier parts of this investigation were carried out 
in the Laboratory of the Biology Department, University of 


326 


Queensland, and to Professor T. Harvey Johnston I wish to 
express my gratefulness for permitting me to perform the 
work there. To him, and to Mr. Henry Tryon, Queensland 
Government Entomologist, I desire to express my obligation 
for the loan of indispensable literature, so difficult to procure 
in Australia. I am also much indebted to the trustees and 
director of the Australian Museum, Sydney, for the per- 
mission granted me to examine important publications under 
their care; and to Mr. W. Rainbow, Museum Librarian, for 
the facilities which he placed at my disposal. Finally, I 
wish to express my sincere thanks to Professor T. Brailsford 
Robertson, of the University of Adelaide, for the many sug- 
gestions and kindly criticisms he has offered me since I have 
known him. 
TECHNIQUE. 

The methods employed here have been fairly simple. For 
the examination of the grosser anatomical processes whole 
mounts stained or unstained, or partial dissections, so far as 
these could be made, have been used. For all the finer 
histological details I have employed sections stained by the 
Heidenhain iron haematoxylin method. TEosin or acid fuchsin 
has been frequently used as a counter-stain. Fixation was 
always made with Bouin’s ‘‘picro-formol’’ mixture. As these 
methods gave very satisfactory results in the majority of 
cases nothing more elaborate was attempted. 


PARTE 


On the Structure and Post-Embryonic Development 
of a Chalcid Wasp, Nasonia. 


A.—TwHeE EXTERNAL FEATURES. 


The eggs of Nasonia, deposited by the female, beneath 
the hard shell of the fly pupa, on to the surface of the delicate 
developing nymph, hatch after a period varying from thirty 
to seventy hours, into small white maggots, about “3 mm. in 
length. These are the larvae in the first instar. 

The larva (fig. 1) is composed of fifteen segments, of 
which the last two can easily be ‘‘telescoped’’ into the one 
preceding them. The last segment is difficult to detect in 
living material. If, however, the larva is placed in a clearing 
solution, which causes considerable shrinking in the cuticle, 
then the segment is unmistakable. 

Of these segments the first two eventually produce the 
head of the adult wasp; the next three develop the thorax, 
while the remaining ten give rise to the abdomen of the 
insect. 

The first segment bears the mouth on its ventral side; 
the last, the anus; but the larva, though it feeds rapidly, is 


327 


quite incapable of defaecating. The first two segments bear 
ventrally a large, powerful, chitinous ‘‘rack,’”’ the tentorium, 
which acts as a support for many of the muscles in the 
anterior region of the animal. The tentorium consists of 
three chitinous bars—really thickenings of the larval cuticle— 
two lateral ones, bent outwards, and an anterior connecting 
bar; while, behind, the structure is supported by a very 
powerful chitinous bar which is formed in the embryo as a 
secretion from a pair of epithelial ingrowths from the walls 
of the second segment. 

The anterior three bars are shed at each moult, and re- 
formed on the new cuticle; they do not reappear in the pupa. 

The mouth is a rather small, transversely elongated, oval 
slit, and is armed on either side by a pair of minute, sharply- 
pointed triangular mandibles, capable of quite active move- 
ment. The head is provided in front with a pair of very 
minute processes, evidently having some sensory function ; 
their nature will be referred to more fully below. 

No other appendages are present. 

Four pairs of spiracles occur; one on the third segment, 
the next on the fifth, the third on the sixth, and the last on 
the seventh. 

The larva feeds rapidly and shows a great increase in 
bulk, an appearance which is accentuated by the imability ot 
the larva to void the intestinal contents. Feeding takes place 


_ by the application of the mouth to a hole torn in the integu- 


ment of the fly nymph by the sharp larval jaws, the food being 
sucked up into the buccal cavity of the larva. The larva 
itself does not appear to move from its orginal place of 
feeding. 

After about thirty hours the larva moults; the second 
instar differs from the first only in its greater size, and in 
the presence now of a-set of nine spiracles. 

The larva undergoes several other moults, but it is very 
difficult to determine their. number, as the various instars 
cannot be recognized by any structural differences. Maud 
Haviland (1920 and 1921) found four instars in two other 
chalcid wasps, species in which differences in the various 
larvae were very obvious. 

After feeding for about three days the larva enters upon 
the “‘resting stage’; food is no longer taken up, and a number 
of remarkable processes begin within the body of the larva. 

Eventually after about a day the larva defaecates, the 
contents of the intestine being voided as minute rounded 
greyish or black pellets; as a result the larva changes from 
a dirty grey to a pure white colour. 

During the next twenty hours—the post-defaecation 


328 


period—the changes commenced in the resting stage continue; 
other changes, which have gone on at a very slow rate during 
larval life, become greatly accentuated. A convenient term- 
ination for this somewhat artificially conceived period is the 
last larval moult, which discloses the pupa (fig. 7). 

Moulting is initiated by a dorsal splitting of the larval 
cuticle. The integument of the pupa is covered with minute 
papillae, which produce a rough surface; and this the pupa 
employs in freeing itself from the larval cuticle. The actual 
ecdysis lasts about an hour, and may best be described as 
taking place by a slow wriggling of the nymph, the larval 
sheath being gradually pushed farther back. 

The liberated pupa has in many respects the appearance 
of the adult insect. The general shape and size of the pupa 
is the same as that of the imago; the antennae, legs, and 
mouth appendages have attained their full length, but are 
thick, “‘fleshy,’’ and ungainly in appearance. In the female 
the ovipositor is quite prominent, lying along the median 
ventral surface of the abdomen. | 

So far, then, as the external features are concerned, the 
most pronounced transformation takes place not in the pupa, 
but in the resting stage and post-defaecation stages of the 
larva. I shall describe first the changes in the external 
appearance of the developing insect as it lies within the 
larval sheath, and then follow the structures, so produced, as 
they continue to develop under the cuticle of the last instar 
—the so-called ‘‘pupal-sheath.’’ This will be followed by an 
examination of the histological processes which bring about 
these remarkable external changes; and finally, the internal 
transformation of the larva will be described. These changes, 
however, must not be regarded as commencing at, or near, 
the time of pupa formation; they have, to'a certain degree, 
been going on during larval life; slowly, indeed, and perhaps 
even spasmodically, but still they have been going on. Some 
time before moulting, however, these changes have become 
accentuated, and others, which have not yet commenced, 
are now initiated; but even these are to be regarded only 
as the result of processes which have gone on in the larva. 

The general shape of the ‘‘living’’ portion of the feeding 
larva is identical with that of the larval cuticle which it has 
secreted, 7.e., it is an elongated ovoid maggot, thick in the 
middle, and gradually tapering at either end. But some 
time before defaecation starts the integument beneath the 
cuticle begins to change its general shape; that of the first 
two segments begins to round itself off, and, before the larva 
moults, has transformed itself into the head of the future 
wasp. A gradual increase or diminution in the size of the 


329 


following three segments gives the thorax the general shape 
that ityhas in the imago; the abdomen shortens considerably, 
rounds itself off, constricts considerably at both ends, while 
at the same time portions of its anterior segments move for- 
wards, and fuse with the thorax to form a compound struc- 
ture, the ‘‘alitrunk,’’ so characteristic of the Hymenoptera. 
The head and its appendages may be considered first. 


The Head and its Appendages. 


The head of the adult wasp is developed from the first 
two segments of the larva; the first segment, to which the 
name oral segment may be applied, develops into the front 
and lower portions of the head, and gives rise also to the 
antennae and labrum. The second segment may be called 
the post-oral segment; from it develop the upper and occipital 
regions of the head, including the ocelli and great eyes, while 
below it produces the maxillae and labium and also the 
mandibles. 

The fact that the first two segments of the larva are 
concerned in the formation of the head can readily be verified 
by following the spiracle of the third segment through the 
metamorphosis, the spiracle remaining as that of the first 
thoracic segment. Already in the late feeding period of the 
larva, the imaginal discs of the head appendages have become 
clearly visible. From the upper portion of the first head 
segment the antennae grow out as short thick processes, 
which, on account of the pressure of the larval cuticle above 
them, are forced to grow downwards (figs. 3, 12). Hach 
antenna has, towards its distal end, a short blunt papilla, 
which fits into the sensory structure on the first segment, 
referred to above. 

Around the mouth, the other head appendages soon 
become prominent; immediately in front of the mouth are 
a pair of quite distinct outgrowths—the rudiments of the 
labrum—which structure is, at this stage, distinctly paired 
(fig. 13). The labrum is generally regarded as a simple, 
unpaired downgrowth from the upper edge of the mouth, but 
in Vasoma its paired condition is quite clear; Patten has 
also figured the labrum as a paired structure in Acilius. (See 
Korschelt and Heider, part i., p. 326, fig. 160). 

The other ‘mouth appendages are developed from the 
second (post-oral) segment; their actual interpretation is, at 
first sight, very confusing, for though they are developed 
from the post-oral segment, some of them take up a position 
actually somewhat in front of the mouth, which is situated 
well within the first segment. The apparent paradox finds 
its explanation in two facts: firstly, the small mandibles of 


330 


the larva, which are merely a chitinous secretion from a part 
of the mass of cells which will later develop the mandibles 
of the adult, are produced from a short mandibular imaginal 
disc, which grows forwards from the second segment and 
terminates close beside the mouth; secondly, in the late larval 
stages there is a considerable shifting forwards of the lower 
surface of the head, the anterior portion of the second seg- 
ment being pushed into the cuticular sheath of the first. (I 
may draw attention here to a fact from which an important 
deduction can be made later, wiz., that the antennae and 
mandibles of the larva, though so absolutely distinct from 
those of the adult, are yet developed in close connection with 
the same group of cells—the antennal and mandibular 
imaginal discs—as produce the corresponding structures in 
the adult wasp.) 

Of the ‘‘post-oral’’? appendages four pairs may be recog- 
nized. ~The most anterior is a pair of short outgrowths, 
which I shall call here the second antennae (fig. 13). Their 
homology will be considered below. Immediately behind these 
arise a pair of long outgrowths, which end close to .the 
larval mandibles—they are the mandibular rudiments; close 
behind these, and nearer the mid-line, is a pair of short 
stout maxillary rudiments; and behind these, and still nearer 
the middle, are the rudiments of second maxillae, quite dis- 
tinctly paired at this stage (fig. 13). 

Of these appendages the mandibles are the largest, and 
I have seen larvae, slightly before defaecation, 7m which each 
1s provided with a palp, which, at this stage, is even longer 
than the mandible itself (fig. 3). I have also observed larvae, 
in the same stage of development, in which no mandibular 
palps were visible. In order to be certain that I was not 
confusing the mandibles with the first maxillae, I examined 
the mouth appendages of defaecating larvae, cut in serial 
sections; under which conditions no error could be made in 
determining the various mouth appendages, and the mandi- 
bular palp could be clearly seen (fig. 48). A mandibular palp 
has not, so far as I am aware, been found hitherto in insects. 
Of special interest, however, is the fact that it does not appear 
to be present in all larvae, its occurrence being perhaps a 
frequent ‘‘abnormality.”’ 

The first maxillae are rather short thick outgrowths at 
this stage, and each has a short palp on its outer side. The 
second maxillae are small, and each has a very distinct palp, 
which twists around the maxilla from below, and embraces 
it distally. At the sides of the second segment are the great 
compound eyes, already differentiating in the late. larval 
period; and in the middle lie the great cerebral ganglia 


(fig. 3). 


“| E 
7 


331 


The post-defaecation period is marked by a continued 
growth of these imaginal rudiments, the completion of the 
process being marked by the last larval moult. 

The head integument bulges outwards, and the biseg- 
mental condition disappears (fig. 12). The head grows, especi- 
ally in height, while above and below the posterior head 
integument grows inwards to form the nearly vertically 


sloping occiput. As a result of these changes, and probably 


also on account of the pressure exerted upon it by the over- 
lying larval cuticle, the head adopts the curious retracted 
attitude so characteristic of the insect. The second segment 
may also be observed, at the end of larval life to be partly 
invaginated into the third. In Calliphora this condition is 
much more pronounced. 

The antennae have meanwhile been growing greatly in 
size. Originally forward outgrowths from the upper region 
of the first segment, the pressure of the larval cuticle soon 
forces them to turn back upon themselves and downwards; 
in the post-defaecation period they grow greatly in length 
and thickness, and at the time of pupation, are in the form 
of two thick appendages, lying ventrally and extending two- 
thirds the distance down the thorax (fig. 7). 

The mouth parts meanwhile continue to grow in size, 
the turning downwards of the head, as already described, 
forcing these into the position in which we see them in the 
imago. Shortly before the larva undergoes its last moult they 
cease to grow, and develop a cuticle on their surface. They 
are now large thick ungainly structures (fig. 14), in no way 
resembling the neat, specialized mouth parts of the imago. 
The labrum is a triangular, irregular flap overhanging the 

mouth; the mandibles are a pair of irregular, ‘‘shapeless’’ 

masses, each bearing its palp, which has now, however, 
creatly degenerated and is little more than a tubercle on 
the mandible. The maxillae are nearly as large as the 
mandibles and project forwards; the palps exceed the maxillae 
in size and are extremely prominent. The labium is quite 
large and from its posterior part project the palps. The 
second antennae have disappeared. 

The remainder of the development of the mouth append- 
ages, during the pupal period, consists of a very pronounced 
shrinking of the structures within the cuticle which they have 
secreted, as a result of which they gradually assume their 
adult shape (fig. 14). 

This process commences a few hours after pupation, 
and within twenty-four hours is practically complete; seg- 
mentation of the appendages has become very marked, and 
bristles are developing on them. The proximal portion of 


- 


332 


the labium (fused mentum and submentum) is almost square 
in shape, and from it spring the medium-sized labial palps. 
The distal portion of the labium (fused endopodites) is slightly 
wider than the proximal portion (fig. 14), and its surface 
is developing a very delicate pubescence. The labial palps 
are rather club-shaped, with a distinct indication of three 
segments; bristles are already clearly visible on them. The 
first maxillae have shrunk to rather short, stylet-like struc- 
tures (fig. 14), and have developed bristles; while the huge 
maxillary palps of the early pupa have shrunk to a pair of 
graceful, four-jointed appendages, on which bristles have also 
begun to develop. © 

The shrinking of the mandibles has been less pro- 
nounced ; each has assumed the shape of a powerful, slightly 
curved jaw, armed distally with three (occasionally with four) 
short blunt teeth; the mandibular palp has entirely dis- 
appeared and its position is indicated, now, only by the 
chitinous tubercle of the pupal sheath. 

The labrum does not undergo any marked changes, except 
a diminution in size. 

As a result of these processes the mouth appendages, in 
the shapes in which we see them in the adult, have been 
produced; chitinisation of this cellular mould, which soon 
ensues, results in the more marked segmentation and the 
hard consistency of the mouth parts, such as we see them in 
the mature wasp. 

The antennae, meanwhile, have been undergoing changes 
parallel to these; at the time of pupation, as we saw, the 
antennae were in the form of two thick, slightly segmented 
appendages lying laterally along the ventral side of the 
thorax. ; 

In the early pupal periods the segmentation becomes 
more distinct, and at the same time shrinking takes place; 
as a result of these processes, the antennae adopt their adult 
appearance after about thirty-six hours; bristles, which are 
first seen some eight hours after pupation, are well marked 
at this stage ; rapid chitinisation of the surface of the antennae 
ensues, resulting eventually in the production of the fully 
developed appendages. 

The antennae of the male and female differ slightly; in 
both sexes there is a long proximal joint, followed bv a 
joint about one-third the length of the first; then come two 
very small joints, followed by nine larger ones, all of about 
the same size. In the female the last three joints are arranged 
so as to form a very distinct club (fig. 11); in the male no 
such modification can be seen. 

The chief point of interest in the development of the 


- mouth appendages is the occurrence of a mandibular palp, . 


| 
| 


333 


probably as an abnormality (since some individuals do not 
appear to show it); the occurrence of this -palp definitely 
proves the homology of the mandible with a metameric 
appendage. . 

The curious nature of the labium is worthy of special 
attention. Its anterior surface, formed as a chitinisation of 
the protoplasmic “‘pubescence’’ already described, is developed 
into a strong rasping-organ. Though present in both sexes 
the ‘‘rasp’’ is more strongly developed in the female. The 
strength and efficiency of the labium is further increased by a 
pair of hard, outwardly diverging chitinous bars lying within 
its distal segment. 

Also to be especially noted is the fact that the head 
appendages first grow in size, and not till the mature size has 
been reached, and even exceeded, does differentiation take 
place. This same fact will be seen also in the development 
of the legs and ovipositor and other appendages; it seems, 
indeed, to be true of the body surface in general (fig. 8): 
first the body becomes moulded, then it begins to undergo 
differentiation producing the various joints, spines, bristles, 
sculpturings, etc., that adorn the insect’s body—first growth 
and arrangement, then differentiation. This fact can be 
demonstrated especially cléarly in the compound eyes (see 
these). The post-defaecation and resting periods are the time 
in which the optic cells adopt their arrangement, in the pupa 
they differentiate. 

Looked at in this light it is possible to regard the pupa 
not merely as an artificially conceived, but as an embryo- 
logically quite distinct phase. Growth occurs in the resting 
and post-defaecation periods; the pupal period is the period 
of differentiation. It should be clearly understood that these 
remarks refer merely to the external characters (integument). 

Meanwhile the great eyes and the ocelli have been devel- 
oping. These structures are merely modifications of the 
integument; already in the resting stage the great eyes are 
clearly recognizable; they grow over a large part of the sides 
of the second segment. In the defaecating larva facets are 
already clearly indicated; these become more distinct as 
development continues. At the end of the larval life the 
_ eyes are very large, and have assumed their typical bulged 
appearance. After thirty-six to forty hours the eyes gradually 
change from a creamy to a pale-reddish colour, which becomes 
bright red a day later. 3 

The ocelli have meantime been developing at the vertex 
of the head, and are seen in the newly formed pupa as three 
prominent rounded tubercles arranged in a triangle. 

After about three and a half days the head gradually 


334 


blackens, and this blackening soon spreads backwards over 
the whole body.; as a result the head is now seen to be marked 
with the sculpturings characteristic of the species. 


The Constitution of the Head. 

Having now described the development of the head, we 
may apply the observations to an attempt to determine the 
metameric constitution of the insect head. , 

Regardless of the actual position which the head append- 
ages have taken up, secondarily, we may enumerate and 
classify them as follows :— 

(a) Pre-oral appendages—antennae, labrum (on first © 

head segment). 

(6) Post-oral appendages—‘‘second antennae,” mandibles, 

first maxillae, labium (on second segment). 

The oral segment gives rise to the face; the post-oral segment 
develops into the occiput, the vertex, and probably into the 
frontal region; from it develop the ocelli and the compound 
eyes. 

‘ This bisegmental condition seems to be very common 
among chalcid wasp larvae. Berlese, for example, in his 
figure of T'apinoma erraticum, actually shows the cerebral 
ganglia lodged in the second ‘segment, while the third 
possesses the first (thoracic) spiracle; the same thing is seen 
in Diachasma, and in the encyrtid wasp Avustralencyrtus, and 
is probably very common, if not universal, among these para- 
sitic hymenoptera. The presence of two head segments is 
especially useful in helping us to determine the homology of the 
insect head. 

The presence of three biramous appendages can be inter- 
preted only in one way, v2z., that three body segments, of 
the primitive annulate-like ancestor of the arthropods, 
gradually moved further and further forward, till eventually 
they became incorporated into the head. This is, however, 
it seems, the usually accepted view; the occurrence of a 
mandibular palp as an abnormality makes the homology more 
certain than ever. What the exact limits of these suppressed 
metameres on the post-oral segment really are, it is not 
possible to say. . 

The oral segment is provided with two pairs of append- 
ages, which have never been observed in a biramous condition 
—the antennae and the labrum. The position of the antennae 
so far forwards, with no appendage in front of them, confirms 
Korschelt and Heider’s view that the antenna of the insect 
is homologous with the crustacean antennule; the small larval 
sensory structures already referred to must, since they are 
formed from the “‘antenna,’’ be likewise homologous with 


i 
2’ 


| 


335 


the Crustacean antennule, or at least with a small portion of 
the antennule. It should be noted, also, that the presence 
of the antenna (antennule) on the oral segment does not pre- 
vent its arising in embryos behind the mouth, a fact which 
Weismann first discovered in Diptera, and which has been 
confirmed by Heider for Hydrophilus, by Patten for Acilius, 
by Nusbaum for Meloe, and by others. 

When, now, we look in the NVasoma larva for the repre- 
sentative of the true antenna of the primitive insect it seems 
that the structure which I have spoken of as the ‘‘second 
antenna’’ must be looked upon as such. It arises from the 
post-oral segment, but actually takes up a pre-oral position, 
and is quite a transient structure. This same post-oral seg- 
ment also gives rise to the eyes and ocelli. 

It seems then that we must regard the insect head as 
composed of at least five segments; the first bears the mouth, 
the labrum, and the ‘‘antennae’’; the second bears the vesti- 
geal true (second) antennae, the ocelli, and eyes; the third 
is represented only by the mandibles; the fourth by the 
maxillae; and the fifth by the labium. 

It is unlikely that the three segments, with biramous 
appendages, should be anything but the first true metameres 
of an annulate worm; the oral segment would then represent 
the procephalic (prestomial) segment of the annulate, and 
the post-oral (with the exception of the three biramous 
appendages) would be the descendant of the cephalic (peri- 
stomial) segment. The presence of the mouth on the second 
segment of Polychaetes cannot be taken as contradicting this 
view, since in the Oligochaetes it is on the first segment. 
The view above expressed receives very strong support from 
the fact that the post-oral segment of Vasonia and the peri- 
stomial segment of the annulate both lodge the cerebral 
ganglia. 

The view above expressed, then, would regard the insect 
head as built up of five annulate segments as follows :— 


Name and No. of 
Segment. 


1. Procephalic seg- 


Represented in 


Represented in 
Imago by— 


Nasonia larva by— 


Oral segment Face, antennae, 
ment Ae | labrum, mouth 
2. Cephalic segment | Apex of head, occi- 
) | put, brain, com- 
| . pound eyes, ocelli 
| Mandibles 


| 
| 
| 


| 
3. First body meta- | 
mere \ 
| 
J 


Post-oral segment 


4. Second body meta- | Maxillae 
mere | 
5. Third body meta- | Labium 
| 


mere 


336 


The extreme view of the constitution of the insect head 
was taken by Savigny (1816) who regarded it as consisting of 
seven segments, corresponding to the antennae, labrum, ocelli, 
great eyes, mandibles, maxillae, and labium; that the 
uniramous antennae and labrum indicate two distinct seg- 
ments, is very improbable; that the eyes and ocelli indicate 
such segments is impossible. Huxley regarded the head as 
constituted, most probably, of six segments. A much better 
conception is that of Lowne (1890), who regards the insect 
head as composed of four segments; his large head capsule — 
(paracephala) lodges the brain, eyes, and antennae and bears 
also two bulbous prominences in front—the (upper) posterior 
cephalocoele, which bears the ocelli, and the (lower) anterior 
cephalocoele. The fact that the posterior cephalocoele bears 
the ocelli shows it to be homologous with the upper part of 
the second segment of Nasonia; the paracephala of Lowne 
are homologous with the remainder of the second (post-oral) 
segment in Nasoma, excluding, of course, the appendages 
which Lowne speaks of collectively as the ‘‘metacephalon.” 
It is the anterior cephalocoele and the neighbourine parts of 
the paracephala which are specially interesting; this region 
bears the antennae, and gives rise in various insects to the 
epistome, the labrum, and the rostrum, and probably the 
mouth ; and there can be no doubt that it is homologous with 
the oral segment of Vasonia. Lowne working with a number 
of insects never found it as a distinct segment, the structure 
having evidently become merged into the paracephala. An 
examination of the Vasonia larva, however, leaves no doubt ~ 
as to its being segmentally distinct. 


The posterior cephalocoele is the Voderkopf of Korschelt 
and Heider. According to Lowne it persists in dragon-flies 
as a bladder-like swelling which lodges the ocelli; its 
persistence in Coleoptera seems to be proved by Lowne’s dis- 
covery of .ocelli as an abnormality in Cuzcindela; in the 
Muscidae a great part of the posterior cephalocoele is with- 
drawn into the rest of the head, as the cerebral vesicles, which 
are evaginated during metamorphosis. 


The only difference, then, between the view of Lowne, 
and that which I have expressed above, is in the bisegmental 
nature of the ‘‘paracephalon.’’ In most insects its existence 
is only a possibility; in Vasonia it is a certainty. In figs. 42 
and 43 of his work on the blow-fly, moreover, Lowne actually 
figures embryos of Calliphora, in which the head consists of 
five segments, and actually appears. to be in a condition 
similar to that of the free larva of Nasonia. His figures 
are taken from Weismann’s great work. 


Y 


337 


The (true) Thorax and its A ppendages. 
The true thorax of the imago is represented by the third, 
fourth, and fifth larval segments, which are all fairly equal 


- to one another in size. Running vertically down each seg- 


ment, on either side, and close behind the spiracle, is a narrow 
streak of integument differing from that which covers the 
remainder of the segment. These six narrow streaks con- 
stitute the imaginal discs of the thorax (fig. 2). 

They are clearly visible through the cuticle of the 
advanced larva, and connected with each are the imaginal 
discs of the thoracic appendages (figs. 2, 5, 6); the first pair 
bear only the first legs; from the second (mesothoracic) seg- 
ment develop the first wings above, and the second legs below; 
the third (metathoracic) disc bears the second wing disc above, 


and the rudiments of the third legs below. 


The wing discs are rather elongated and can be seen to be 
enveloped in a distinct sac. The leg discs are much shorter 
than those of the wings; and the sacs in which they are 
carried are very distinct, each bearing a small opening on to 
the surface of the integument, below the larval cuticle. 
Neither in the wing, nor in the legs, could I detect any 
indication of a biramous structure. 

During the resting stage of the larva the imaginal discs 
begin to grow rapidly; the integumentary discs spread out in 
all directions, the first two, especially the second, rapidly; 


the last very slowly. The discs of the appendages soon grow 


out of their sacs; already in the larva at the time it defaecates 

the legs have protruded so far that they begin to bend upon 

themselves beneath the larval cuticle, and we see the earliest 

indication of segmentation. The wing discs, on the other 

ee ade downwards as two large sacs, and do not bend 
@. 3). 

About ten hours after defaecation the discs have, to a 
large extent, assumed their imaginal shape and size; not till 
about the time of pupation, however, as will be seen later 
when we examine the histological structure of the developing 
discs, is the process of encroaching quite complete. The first 


thoracic disc is now seen to have projected forwards to form 
a hood over the upper part of the head; the second disc 
_ has far outstripped the other two, and, growing right under 


the cuticle of the first thoracic segment of the larva soon 
assumes its imaginal dimensions; from it about three-quarters 
of the thorax develops; the metathoracic disc scarcely 
lengthens at all, and persists as a small ring behind the great 
mesothoracic segment. 

The legs and wings have meantime been extending, and 
already in the larva a few hours after if has defaecated the 


338 


process of segmentation of the legs has proceeded far. In 
the larva ten hours after defaecation a coxa is distinctly 
visible behind the femur; the region where the leg showed its 
original bending marks the beginning of the tibia; the tarsus 
is also clearly visible; on the third tarsus at least four seg- 
ments have been produced, on the other tarsi joints are just 
forming. No trochanter is visible yet. 

The wings have meanwhile grown in size. The first wings 
are now in the form of two great hollow sac-like pockets, on 
either side of the mesothoracic segments; the hind wings are 
much smaller. 

In the larva a few hours later the legs have grown so 
long that they are found beneath the thorax, and their distal 
ends begin to grow backwards; the wings continue to grow in — 
length, and likewise become forced backwards. 

The proximal wide ‘‘mouth’’ of the wings contracts more 
and more, and the great sac-like structures transform, in the 
late larva, into others having the shape more nearly of the 
wings of the adult. Rapid growth of the legs continues, so 
that just before the end of larval life the first leg has grown 
backwards nearly to the end of the thorax; the second about 
one-third the distance down the abdomen; the third about 
one-quarter the length of the abdomen from the end. The 
" wings also, especially the first wings, have become very large 
and have enveloped the sides of the thorax. 

All the segments of the legs, except the trochanters, 
are cleary seen; but the legs themselves are thick fleshy struc- 
tures, resembling only in a general way the legs of the adult 
(fig. 16). The same thing has been described above in the 
mouth-appendages. 

At an early stage in their formation as distinct append- 
ages, 2.e., in the resting larval period, tracheoles began to 
extend into the wings and legs. Each leg is provided with 
a single long tracheole occasionally branching into two parts 
distally. The wings, on the other hand, are well provided 
with tracheoles. Their actual structure and their history 
within the wings will be described later; it will suffice to 
refer here merely to their general disposition within the 
wings. 

Running along the lower (anterior) border of the wing 
are a pair of tracheoles (figs. 44, 65), one of which is bifur- 
cated distally. They appear to communicate in the proximal 
portion of the wings. Passing down the middle of the wing 
are a number of tracheoles, which appear also to be branches 
of a single large tracheole at the base of the wings, this large 
tracheole giving off a pair of smaller vessels, each of which 
bifurcates in about the middle of the wing. One of these 


| 
4 


339 


small vessels passes to the end of the wing and there turns 
upon itself and runs forwards again. These tracheoles, as 
well as those of the legs, are all markedly twisted and 
irregular. ’ 

Meanwhile the thoracic segments have been undergoing 
further development. The second segment becomes somewhat 
convex; the first segment grows downwards, and instead of 
overlying the rear of the head, now comes to assume its 
proper position as a shield over the neck and front of the 
thorax. A mechanical explanation of this will be given later 
(see Muscular System). 

The metathoracic segment retains its insignificant size. 

Having arrived at their maximum size, the thoracic seg- 
ments and their appendages form a cuticle. This process, 
which is coincident with cuticle formation over the rest of 
the body, is quickly followed by the pupal moult. 

The thorax, which has now attained its general adult 
shape, begins to undergo changes parallel to those that go on 
in the head, 7.e., ridges, grooves, tubercles, bosses or depres- 
sions, etc., are developed on its surface in the positions in 
which we see them in the imago. This process takes place 
on the first day of pupal life, and is soon followed by chitinisa- 
tion. Already in the thirty-six hour pupa this has pro- 
ceeded considerably, and the only changes which take vlace 
during the remainder of the pupal life consist in a thickening 
of this chitinous coat, accompanied by a general blackeiing, 
following close on the blackening of the head. 

The legs, meanwhile, undergo continued ‘‘differentia- 
tion’’; they shrink greatly within their cuticle, and the 
segments become more clearly marked (fig. 16). Already at 
about six hours after pupation the shrinking has permitted 
the growth of (protoplasmic) bristles on the surface of the 


legs; soft claws and spines are soon seen, and the trochanters 


are clearly visible some twenty-four hours after pupation. 
The view of Lowne that they are really part of the femur, 
and that they do not represent a distinct segment, com- 
parable, for example, to the coxa, or tibia, seems justified 
by their very late appearance in the pupa, the true segments 


being clearly visible even in late larval stages. Some twenty- 


four hours after pupation the legs have practically assumed 


_ the external appearance of those of the imago. This is fol- 


lowed by the secretion of chitin, at first slow, later rapid, so 


_ that at the end of two and a half days the legs of the pupa 


are (to external appearances) identical with those of the 
adult wasp. 

_ The wings, also, have continued to develop during this 
time. First a considerable shrinking takes place, so that the 


» 4) 
340 ‘i 
wing occupies, in the twenty-four hour pupa, an area about 
three-quarters the size of the wing of the newly formed pupa 
(fig. 44). Already in the pupa a few hours old, the upper 
and lower pairs of veins of the fore wing are seen to he 
each within a broad clear space, extending from the base of 
the wing to a distance about one-fifth the length of the wing 
from the end. In the twenty-four hour pupa these clear areas 
have become much more distinct. In the hind wings, so far as 
I could observe, only a single such clear area is formed. The © 
wings meanwhile have assumed, more nearly, their adult 
shape, showing now a very slender proximal region, and 
developing, at the same time, each a small basal structure 
provided with a number of irregular prominences and depres- 
sions (fig. 9), which articulate with, or into which fit, other 
depressions and projections from the sides of the thorax. 

The wing now assumes a remarkable appearance; instead 
of remaining as a smooth fleshy structure, its surface begins 
to undergo, in the thirty-six hour pupa, a very pronounced 
folding (fig. 37); at the same time hairs—the fine pubescence 
of the adult wing—-as well as bristles, and the hooks of. the 
hind wings begin to appear on the surface. The folding is 
soon complete, and the whole structure now begins to chitinise. 

The chitin on the hooks of the hind wings becomes fairly 
thick; elsewhere, however, the chitin remains thin, and 
closely follows the contours of the wrinkled surface of the pupal 
wing. 
The anterior clear space of the fore wing, and that of 
the hind wing become brown in colour; they form the nervures 
of the adult wings; the bifurcated clear areas on the rear 
half of the first wing do not change colour, and remain as 
colourless ‘‘pseudo-nervures,’’ so characteristic of the wings 
of many chalcid wasps. ! 

On emerging from the pupa the wings of the wasp soon 
straighten out. Not till now can we actually estimate the 
extent to which folding of the wing epithelium has taken 
place; for so pronounced has been the folding within the 
limited space afforded by the pupal cuticle covering the wings, 
that these, on expanding fully, attain, in the course of a few 
minutes, an area sixteen times that of the pupal wing. 

At the rear of the fore wing a slight turning over of the 
wing chitin forms the only structure on which the great 
hooks of the hind wing can possibly find a grip. 

The actual cellular processes which underlie this remark- 
able development of the wings will be described later (see 
page 359). 

One point seems to be worthy of special attention here. 
The fact that the clear spaces (developing nervures) of the 


341 


pupal wing may at times be quite devoid of tracheoles, while 
at other times they may lodge two tracheoles, if not more, 
seems to show that the nervures have no relation whatever to 
these respiratory tubes, and that the latter have grown into 
them merely because they represent a path of diminished 
resistance to growth. It seems to follow, also, that attempts 
to arrive at any conclusions as to the phylogeny of families 
and genera (in the chalcid wasps, at any rate) on the basis 
of pupal wing structure are fallacious, unless special distinc- 
tions are drawn between the wing nervures and the tracheoles 
which they may contain. 


The Abdomen and its Appendages. 


The abdomen of the wasp is built up from the last ten 
segments of the larva, and in its general features the develop- 
ment is identical with that of the thorax, 7.e., the narrow 
imaginal discs of each segment grow outwards beneath the 
cuticle of the larva and assume the general shape of the 
abdomen of the wasp. The imaginal discs, moreover, are 
similar in appearance to those of the thorax (fig. 2), being 
in the form of narrow strips of tissue, running vertically 
down each segment close behind the spiracle (in those seg- 
ments which possess one). The imaginal disc of the last seg- 
ment occupies the whole of its lower lateral regions. 

The general shape assumed by the abdomen is ovoidal; 
but in this the first two segments do not co-operate; on the 
other hand, a remarkable migration takes place here, and the 
whole of the first abdominal segment, and the upper half of 
the second, become merged in with the thoracic segments, to 
form the middle region of the insect, the hymenopteran 


“alitrunk,” while the lower part of the second abdominal 


oe 
oe 
v 
= 


segment forms the petiole, which in the adult wasp connects 
the “‘abdomen”’ with the alitrunk, and articulates with the 
upper part of the second segment, with which, therefore, it 
always remains in fairly close intimacy. 

It is in the larva about twelve hours after defaecation 
that this process of migration is first indicated. At this stage 
a horizontal splitting is seen in the second abdominal seg- 
ment, and shortly after, the first segment, and the upper 
half of the second, begin to move forwards, while the lower 
portion of the second retains its position, and eventually forms 
the petiole. At the time when the larva moults, these two 
segments have distinctly left the remainder of the abdomen, 
and have produced the ‘‘alitrunk’’ (fig. 7). At this time, 
also, the abdominal discs have completely encircled the body, 
and the lower portion of the second abdominal disc has so 
constricted as to give it the form of the petiole; already in 


342 


the larva fourteen hours after defaecation the petiole is clearly 
seen. As on the rest of the body, the attainment of adult 
proportions is rapidly followed by the secretion of a delicate 
cuticle, after which the larva moults. 

The changes which occur during pupal life in the 
abdomen are quite parallel to those occurring on the 
remainder of the body. A certain amount of shrinking 
takes place some twelve hours after pupation; bristles form 
on various parts of the body; the small sculpturings of the 
wasp’s body are moulded on the soft epithelium of the pupa, 
and then the whole abdomen undergoes chitinisation. The 
blackening of the abdomen takes place on the fourth day, 
soon after that of the head and thorax, 7.e., the wasp blackens 
from before backwaids. 

The two abdominal segments which have become merged 
into the alitrunk undergo but slight changes during pupal 
life. Just as the first thoracic segment grows downwards 
to form the front wall of the thorax, so the two migrated 
abdominal segments also grow downwards to form the rear 
wall of the ‘‘alitrunk.’’ The two processes take place at the 
same time, and are to be explained, I believe, as the result 
of a pull, exerted upon them by the contraction of the great 
longitudinal thoracic muscles, which pass horizontally from 
the one to the other (see below, p. 426). This pulling down- 
wards results in the more distinct separation of the alitrunk 
from the rest of the abdomen. ‘The chitin in this region 
becomes extraordinarily thick, and the whole surface under- 
goes remarkable sculpturing. In the two-day pupa, the upper 
part of the second abdominal segment is still clearly visible 
as a small square segment embedded in the one preceding it; 
it is about equal in size to the petiole, with which it articu- 
lates. As chitinisation advances, however, it becomes more 
and more difficult to detect. 

That the alitrunk contains the first abdominal segment is, 
of course, well known. But that the upper portion of the 
second segment is also incorporated in the alitrunk does not 
seem to have been recognized hitherto. Thus Sharp (1895) 
writes :—‘‘The structure of the posterior part of the alitrunk 
has given rise to an anatomical discussion that has extended 
over three-quarters of a century, with the result that it is 
now clear that the posterior part of what appears to be thorax 
in Hymenoptera is composed of the abdominal segment. This 
part has been called ‘Latreille’s segment,’ the ‘median seg- 
ment,’ and the ‘propodeum.’” . . . ‘‘Although the true 
first segment of the abdomen is detached from its normal 
position and added to the thorax, yet the term abdomen is 
conveniently restricted to the part that commences with the 
true second segment’’ (part 1, p. 492). 


343 


To what extent this bisegmental condition of the pro- 
podeum is found in Hymenoptera generally I am unable to 
say. In Nasonia there can be no doubt of it, but to deter- 
mine its constitution in other groups would necessitate an 
embryological examination of these, the study of mature 
material being generally useless. 

It remains only to describe the segments of the rest of 
the abdomen. The third segment is large and conical and 
overlaps the second, which is rather shorter; the next is very 
long, and the following two rather shorter. The last four 
segments are small and together form the posterior quarter 
of the abdomen (fig. 4). 

Chitinisation has taken place so as to produce distinct 
terga and sterna. 

The sterna of the seventh and eighth segments do not 
meet below, the body being protected here by the sternum 
of the ninth segment. The tenth segment is partly invaginated 
into the preceding one and is represented by a terminal plate 
which bears a long horizontal slit, the anus (figs. 21, 26). 
Surrounding this plate are other podical plates, but whether 
these are formed from the last, or from the antepenultimate 
segment, I am not definitely able to say. The last segment is 
interesting in that it bears in the female a. pair of processes, 
which grow out some ten hours before putation ; in the newly 
formed pupa they are in the form of short blunt appendages 
a little longer than broad, quite prominent in ventral view 
ng 21’). 

In ie early pupa the usual contraction takes place, and 
they remain as short conical papillae at either side of, and 
just below the anus; each is covered with long bristles, 
developed early in pupal life. These are the tactile hairs, 
and the structure serves as a delicate sense-organ for the 
female in the examination of the surface of the fly-pupa for 
a suitable spot to pierce with her ovipositor. 

The modifications which these structures undergo in the 


male will be described in connection with the development of 


the male copulatory organs; histological details will be given 
in connection with the description of the integument. 


In the female, meantime, the ovipositor has been develop- 
ing. This is represented in the feeding larva in its last instar 
by three pairs of imaginal discs, situated in the twelfth, 
thirteenth, and fourteenth segments, and identical in appear- 
ance with the anlagen of the legs (fig. 2); there is no evidence 
of a biramous structure. Some time before defaecation these 
appendages grow out above the surface; the upper pair grow 
backwards along the ventral side of the insect as two hollow 


344 


appendages, lying parallel to each other (fig. 20); the second 
appendages similarly grow backwards ventral to these, also 
as two hollow appendages, but more closely applied to each 
other; the last pair, on the other hand, extend mainly for- 
wards, and serve as a lateral protection for the other four 
appendages which the former partly enclose. Growth is rapid, 
and the posterior appendages assume the function more and 
more of a protecting sheath for the other two pairs. Already 
in the defaecating larva the second pair are closely adhering; 
in the larva eighteen hours later the two have almost formed 
one tube, and by the time the larva moults, there can no 
longer be any doubt as to the occurrence of a distinct cavity 
in this structure. This cavity, however, is not formed by a 
fusion of those of the two appendages; on the other hand, it 
is the result of an incomplete fusion of the walls of the two 
appendages, due to the invagination of the inner half of each 
into its outer half. The anterior pair of appendages mean- 
while grow in thickness, and tend to fill the available space 
enclosed by the last pair. 

The first pair of segments meanwhile give off each an 
anterior offshoot which grows upwards and curves backwards 
into the abdomen of the wasp (fig. 21); from the end of this 
outgrowth, a second portion grows downwards and forwards. 
This process seems to be complete at the time of pupation. 
Already in the larva, eighteen hours after defaecation, a 
distinct chitinous cuticle has been formed around the external 
parts of the ovipositor, and the organ is now prepared for 
the moult. At the time of pupation, then, the ovipositor 
has assumed its general adult appearance, but as with all the 
other appendages, the structures are merely moulds for the 
adult organs—they have attained their required sizes; they 
have now to differentiate. 

This takes place in the early pupal period. The first 
pair of appendages shrink, and tend to enclose the second 
fused appendage, which has also shrunken; the posterior pair 
of appendages similarly shrink, and remain as somewhat 
flattened thick sheaths, protecting the others. (fig. 22). 

The aperture of the female sexual ducts is a wide trans- 
verse slit-like structure, and this opens into the cavity enclosed 
by the first and second appendages. It will be referred to 
more fully in connection with the female sexual organs. The 
second appendage becomes serrated distally and projects 
slightly beyond the tips of the first pair. The whole struc- 
ture, in the two-day pupa, then undergoes chitinisation, to 
form the ovipositor of the adult. 

This complex organ consists, then, of a protecting sheath 
(third appendage) which encloses the actual egg-depositing 


ee, 


345 


tube. The latter consists of a slender but very stout rod, 
serrated distally (fig. 22), whose function is to bore through 
the fly-pupa prior to oviposition, the eggs entering the hole 
through a tube formed between this rod and the pair of first 
appendages which partly surround it. The proximal portion 
of these first appendages has grown in a strong curve, as 
described above, into the abdomen of the wasp ; a second piece 
growing forwards and downwards from the end of this has 
also been described above. These structures likewise chitinise 
and produce an exceedingly efficient system of phragmas. 
The insect has, as it were, taken full advantage of this, and 
a great group of muscles has developed, whose function is to 
move and hold the ovipositor while the latter is functioning. 
These muscles are shown in fig..22. One group radiates out 
from the base of the second appendages, and is inserted on 
to the first portion of the phragma. . Other muscles are 
inserted into the descending portion of the phragma; others, 
again, are attached to the base of the ovipositor. A system of 
smaller phragmas is also developed on the ventral body wall 
to give firmer attachment to the “‘origin’’ of these muscles. 
The figure, however, will make this elaborate system of muscles 
clearer than any verbal decription can. 

The action of the ovipositor is now obvious; a pull by the 
muscles of the great phragmas will immediately swing the 
ovipositor forward out of its sheath (third appendage) into a 
vertical position, and the prolonged contraction of these and 
other muscles holds the ovipositor very rigidly for several 
minutes, during which the upward and downward movement 
of the abdomen causes the rigidly fixed ovipositor to bore its 
way through the hard sheath of the unfortunate fly-pupa. 

It is worth noting here, that during the development of 
the abdominal imaginal discs, the eleventh body segment (sixth 
abdominal) grows backwards a considerable distance along 
the ventral body wall and overlaps more than half the anterior 
portion of the ovipositor. During oviposition, consequently, 
when the very flexible abdominal segments are subjected to 
considerable strain, the ovipositor pushes these overlaps for- 
wards and has, then, the appearance of arising from a 
pyramidal structure on the ventral side of the body. No such 
structure is, of course, normally present. 

The histological changes which underlie this development 
will be referred to under ‘the integument.”’ 

_ The devolpment of the ovipositor of Locusta has been 
described by Dewitz (1875). He describes an anal segment 
bearing a pair of appendages (cerci) homologous probably with 
the sensory papillae of Vasonia. The ovipositor is developed, 
according to Dewitz, from the three preceding segments, one 

K . 


346 


of which is formed late in life. The only apparent difference 
between the structure seen in Locusta and that found in 
Nasonia lies in the fact that the two second appendages do not 
unite (in Locusta) to form a boring organ, and that the third 
appendages do not merely act as a protecting sheath for the 
ovipositor, but actually enter into its formation. | 

It is necessary to describe next the formation of the 
copulatory organs in the male. 


So far as I could observe, no rudiments corresponding to 
those of the female copulatory organs are present in the male 
larvae, and no special differentiation of external male organs 
takes place till very late in larval life. This is intimately 
connected with certain changes in the last four abdominal 
segments, which become so disposed as to allow of the 
eversion of the penis. © 

Shortly after the defaecation period the tenth and ninth 
imaginal discs of the abdomen have grown so as to assume 4 
position at the rear of the animal and at the same time to take 
only a very small part in the formation of the lateral, ventral, 
or dorsal walls of the larva. This is brought about by the 
fact that the growths of these two abdominal imaginal discs are 
not very extensive; and that they actually become partly invag- 
inated into the eighth abdominal segment, which grows much 
faster than they do. This is clearly seen in the section shown 
in figs. 25 and 27. The tenth (terminal) segment is quite 
small, and bears the anus; ventrally it is provided with a pair 
of appendages, which lie in close contact with the ninth 
segment, which segment is partly invaginated into the eighth. 
This invagination is accompanied by a marked cell-prolifera- 
tion in the integument of the invaginated portion (fig. 25), 
and is already clearly seen in the defaecating larva; during 
pupal life it develops into the penis. 

The segments then chitinise and are found, in this con- 
dition, in the early pupa. It is especially worthy of notice 
that already at this stage the two appendages of the tenth 
(terminal) abdominal segment have applied themselves very 
closely to the sternal portion of the ninth (fig. 27); indeed, 
while the segments of the cuticle of the pupa are, in other 
respects, an exact representation of the shape of the epidermis 
which has secreted them, yet the appendages of the tenth 
segment form with the sternal portion of the ninth a cuti- 
cular covering which is common to them both; an examina- 
tion of the epidermal components of this compound cuticular 
segment, however, reveals its true nature (fig. 27). In the 
pupa, shortly after its formation, a curious change now takes 
place, which results in the formation of the penis. The cells 


— 


347 


of the sternal region of the ninth imaginal disc continue to 


- proliferate rapidly, and grow inward, into the eighth seg- 


ment; the extension is very rapid, so rapid, indeed, that after 
about six hours the invagination has extended forwards as 
far as the posterior border of the sixth segment; the process 
continues, and does not cease till the invagination has. ex- 
tended, along the mid-ventral region, well into the fifth 
abdominal segment. 


Early in the process of invagination a , cavity is developed 


in the mass of cells; and at the terminal (anterior) end this 


cavity dilates, to form a small sac, the vesicula seminalis 
(fig. 27), which is, therefore, formed from deeply invaginated 
epidermal cells, and is in no way to be regarded as a meso- 
dermal structure. In the six-hour pupa, though already 
clearly defined, it has, nevertheless, not attained a very 
pronounced size; but some twelve hours later, it is quite a 
prominent bulbous dilatation at the anterior end of the penis. 


The penis is thus a structure composed of the sternal 


portion of the ninth segment, and the appendages of the 


tenth, and the whole organ is produced simply by a massive 
ingrowth of cells of the ninth segment, forwards, along the 
ventral body wall. 


Already in the earliest pupae the transference of the 
appendages of the tenth segment to the sternum of the ninth 
is clearly visible, but it is not till well within the second day 
that the distinct development of a joint separating the two 
is evident. Early in the pupa the two appendages unite to 
form a simple tube, but exactly how this takes place I have not 
been able to observe. 

The penis is, then, a simple tube, consisting of two por- 
tions, a proximal, representing the sternum of the ninth 
segment, and a distal shorter portion, developed from the 
appendage of the tenth segment. The distal segment is seen, 
in the pupa, to be invaginated into the ninth; and both the 
segments are provided with a pair of long tendons, which serve 
to withdraw the distal joint into the one preceding it, and, 
finally, the whole structure into the abdomen. In this con- 
dition the organ is seen during later pupal life, and the ventral 
termination of the abdomen of the male, though really so 
totally different from that of the female, has, nevertheless, a 
curious resemblance to it. This is due to the fact that develop- 
ment of the male copulatory organ is mainly a process taking 
place within the abdomen, after the pupa has been formed. 

This internal development of the tenth and ninth seg- 
ments is accompanied by a number of changes in other 
abdominal segments, which result, in part, in the formation 


K2 


4 


of the accessory copulatory organs of the ninth segment, or 
in the modifications of the segments to aid in the eversion of - 
the penis. In the larva shortly before pupation, the ninth 
segment develops terminally a pair of great ‘‘beak-like” clasp- 
ing forceps, which have a very important accessory copulatory 
function, while the sternal regions of the seventh and eighth 
segments, which have not kept pace with the extension of 
the tergal region of these segments, become pushed forwards 
and are partly invaginated as the penis develops (fig. 26). 

Shortly after the penis adopts its adult proportions 
muscles become developed within it. 

The histological processes underlying the changes will be ~ 
dealt with later in connection with the development of the 
integument. 

The general blackening of the cuticle of the nupa com- 
mences some three and a half days (in summer) after the last 
larval moult, and is complete about twelve hours later; the 
wasp remains enclosed in the pupal sheath for twelve to 
twenty-four hours longer, and then, splitting the thin sheath 
which imprisons it, escapes. 


348 


B.—TueE IntTEGUMENT (Histological Development). 


In the newly hatched larva the integument has reached 
a state of development, which it retains with but small changes 
throughout the feeding period. The ectoderm consists, for its 
greater part, of a single layer of cells which are of two kinds; 
there are the large cells, less numerous than the other type, 
but occupying a greater part of the integument—the true 
“larval-cells ;’’ and, secondly, there are the narrow strips of 
integument consisting entirely of smaller more embryonic 
cells—the centres from which the imago will later develop— 
the imaginal discs of the integument; indeed, at this early 
period the rudiments of the wings, legs, mouth appendages, 
antennae, and even of the eyes are clearly recognizable, while 
the areas from which the general body surface of the imago 
will later develop are very prominent. 

It is to the development of the general body integument 
that we will first give our attention ; this will be followed by 
the description of the formation of the legs, wings, antennae, 
and mouth appendages; and, finally, the most astonishing of 
ali the integumental changes, the development of the 
compound eyes and ocelli will be described. 

In the newly hatched larva the cells of the imaginal discs 
of the general body surface are small, short, and columnar, 
and closely packed side by side (fig. 10); their protoplasm is 
clear, and their nuclei are very large. As the larva grows 


349 


cell division takes place, so that in the larva, at the time it 
defaecates, the integumental imaginal discs contain about four 
times the number of cells that we see in the newly hatched 
larva. Whether these cell divisions occur during the moulting 
period, or whether they occur gradually throughout larval 
life I am unable to say. Even this extensive multiplication 
is not sufficient, however, with absence of actual cell growth, 
to enable the cells of the imaginal discs to maintain their early 
appearance and at the same time retain their function of 
forming an unbroken body layer, such as is necessary in the 
secretion of a new cuticle in the period just preceding a moult. 
The difficulty is overcome by the cells gradually assuming a 
curious shape; their outer ends develop into thin flat discs. 
their inner ends containing the nucleus become long and 
narrow. Thus, while the outer portions of the imaginal 
embryonic cells combine to present an unbroken surface—a 
tiue pavement epithelium—the inner ends, which give the 
predominating appearance to the disc, are long and narrow 
and separated by wide spaces. The large specialized larval 
cells, on the other hand, do not undergo these changes; on the 
contrary, they retain their early shape and number, and the 
only visible change which they undergo is a great increase in 
size, an increase approximately proportional to the growth of 
the larva as a whole. 

These large larval celis of the imaginal discs co-operate 
during the feeding period to form the various cuticles. 
When newly formed, the cuticle, which is simply a direct 
secretion from the ectodermal cells, embraces these very 
closely; gradually it loosens itself, when the integument 
begins to secrete a second cuticle, inside the first, the two 
cuticles being very clearly visible in sections through the 
integument (figs. 47, 55). 

At the time when the larva begins to defaecate the 
ectodermal cells begin to enter upon a period of profound 
changes. The nuclei have become very large—indeed, their 
growth appears to have kept pace with that of the whole cell 
—but the chromatic contents appear curiously disorganized. 
Each contains a relatively gigantic nucleolus. Then follows a 
period of cytoplasmic disintegration. The entire cell contents 
break up into numerous minute globules (figs. 28, 29) which 
have a curious resemblance to nucleated cells, each consisting 
of a clear outer zone, and containing a heavily staining body. 
They are, however, purely disintegration products of the 
large cells and are not to be confused with the leucocytes, 
which are much larger than these globules; their curious 
construction is probably due to some obscure physical con- 
dition of the disintegration products of the cell. These 


a - 
4 
‘ 
J ' 


350 


globules break loose from the cell, and are apparently 
dissolved in the blood. 

The disappearance of the larval cells 1s not, however, 
entirely one of chemical disintegration; definite phagocytosis 
of the cells also occurs, but its action is quite secondary to 
that of the chemical disintegration. Indeed, it is improbable 
that there are at this stage enough leucocytes in the blood of 
Nasonia to bring about the destruction of the larval integu- 
ment—not to mention the destruction of other larval organs. 
‘That the process does occur, however, is quite certain; 


leucocytes may frequently be seen lying upon or within the 


disintegrated cells, and filled, at times, with degeneration 
globules, which they have recently ingested (fig. 31). They 
are the Kornchenkugeln of Weismann. 

Sometimes the cell contents do not break up into these 
minute ‘‘pseudo-nucleated’’ globules, but the whole mass 
undergoes granular degeneration and produces a large ball 
(fig. 30), which, after lying for some time within the cell 
membrane, breaks loose, and tumbles into the general body 
cavity; a few globules generally remain within the cell mem- 
branes, which now appear as irregular, empty hulks, below 
the developing imaginal integument. Sometimes, again, the 
cell contents may degenerate into large hyaline spheres, about 
the size of leucocytes, each containing several heavily-staining 
granules (fig. 28). 

In the body cavity the big granular spheres are fallen 
upon by the leucocytes, and by the intervention of these, and 
to a certain extent, apparently, by a process of solution, they 
gradually disappear. This type of cell disintegration is especi- 
ally clearly seen in the larva about sixteen hours after 
defaecation. 

The process of integument destruction lasts nearly a 
whole day; the cell contents first disappear, leaving only a 
thin cell membrane, which, in turn, eventually disintegrates. 

Accompanying these changes in the integument there is 
a total renovation of the underlying somatopleure. The larval 
somatopleural cells are greatly overgrown, and present a large 
nucleolus. Smaller embryonic cells, which have evidently 
lain dormant within the somatopleure, begin to proliferate 
during the period just preceding defaecation, and growing at 
the expense of the larval cells which they absorb, finally 
redevelop into a new somatopleure. The splanchnopleure, 
covering the internal organs, undergoes similar changes. 

Meanwhile the imaginal discs have become active, and, while 
the cells undergo further multiplication, begin to encroach 
upon the places occupied by the disintegrated larval cells, 
eventually replacing these entirely. The discs grow out in all 


es 


351 


directions; the outer, most actively migrating cells often 
show amoeboid processes (fig. 29), and it may be by this 
method of locomotion that the cells advance. Sometimes they 
grow right over the dead larval cells (fig. 28); at other times 
they seem to be unable to cross them, and have to await the 
complete destruction of the larval integumental cells before 
they can advance further (fig. 32). 

During the last hours of larval life, therefore, the integu- 
ment consists of areas of proliferation, the cells of which, 
growing outwards, are actively engaged in replacing the dis- 
integrated larval cells, or awaiting the total destruction of 
these. 

Eventually the imaginal discs of the integument meet, 
a cuticle is secreted and the larval moults, disclosing the pupa. 

At times leucocytes, having disposed of the remains of 
the larval tissues, are seen crammed with larval débris lying 
among the proliferating imaginal cells, and evidently provid- 
ing, by their disintegration, nourishment for the surrounding 
cells (fig. 31). Sometimes, also, numerous of the large hyaline 
degeneration globules are seen in similar situations (fig. 31). 
The integumental cells, unable to extend further, now 
begin to undergo structural changes; at first spindle-shaped 
(fig. 30), they soon begin to change their general form; in 
some parts of the integument, especially that of the abdomen,’ 
the cells are small and cubical, their outer surfaces very 
regular; the chitin secreted from them in this region is quite 
smooth. Along the dorsolateral regions of the pupa the cells 
are generally rather elongate. In the antero-dorsal region 
of the thorax (especially in the region of the future pronotum) 
this condition is especially clearly seen. Here the outer ends 
_ of the cells develop broad swellings, giving the cells a hammer- 

like appearance; from these swellings the thick chitin in this 
region becomes secreted. A somewhat similar condition is 
seen right at the posterior end of the abdomen. The ectoderm 
of the propodeum and metathorax is especially remarkable, 
being in the form of a great accumulation of ectodermal cells, 
several layers deep, all crushed together, and thus accounting 
_for the contraction which external features show has gone 
on in this region. 

The secretion of cuticle now goes on very rapidly, and 
about four hours after pupation, forms a layer nearly as 
thick as that of cells which are secreting it. The cells from 
which the cuticle is being secreted, moreover, do not, as a 
rule, present a perfectly smooth surface, but become so dis- 
posed as to form a mould on which the cuticle of the imago 
can shape itself, and the various depressions and bosses, and 
other sculpturing with which the imago is ornamented, as well 


352 


as the hard bristles and claws, and even the delicate hairs 
(pubescence) are to be regarded simply as chitinisations on 
the surface of these cells, or on parts of them. The ‘‘spiral’’ 
thread of the tracheae, as will be described later, is similarly 
merely a chitinisation of a previous protoplasmic ‘‘mould.” 
The small sculpturings on the body surface generally require 
only a single cell to act as a mould for them; this is very 
clearly seen, for example, on the head (fig. 33), the dorsal 
part of which shows forwardly projecting scale-like bosses, 
while on the antero-ventral part these project upwards; and 
the cells, in early pupae, can be distinctly seen, one under each 
boss, and all disposed in such a way as to present a ‘‘scale- 
like’’ appearance similar to that of the imaginal cuticle which 
they are secreting. The larger sculpturings, as well as such 
structures as claws and large spines on the legs, are, as a 
rule, moulded upon a number of cells (fig. 34). 

Bristles, on the other hand, are unicellular structures. 
Their formation can be especially clearly seen on the ovi- 
positor and posterior extremity of the insect. The ectodermal 
cells begin to elongate and develop a point at their free ends ; 
the elongation becomes more and more marked till the cell 
assumes the slender form of the bristle as we see it in the 
imago. Then it begins to chitinise (fig 35). The insertion of 
such a bristle on the cuticle of the imago is always strengthened 
by a small ring-like supporting structure (fig. 35), and the 
protoplasmic mould even of this support can, if the hair and 
cell is observed at the right moment, be clearly seen. 

The development of minute hairs (pubescence) is especi- 
ally curious. The process can be clearly observed on the 
second maxillae (labium). Here the ectodermal cells develop 
a number of long delicate processes, giving the cells a curiously 
frayed appearance at their terminations (fig. 24). Each of these 
processes then chitinises, to form a single hair. A single cell 
therefore acts as a mould for a number of minute hairs and the 
co-operation of a number of such cells produces the rasp-like 
pubescent structure which one finds on the ‘“‘tongue-like’’ 
labium of the adult Wasona. The chitinisation of the 
epidermal cells continues throughout pupal life, and the pro- 
cess does not cease till the whole of the cells have been 
converted into chitin. A cellular ectoderm is, therefore, 
absent in Vasonia, except in the region of the great eye. 

The extraordinary accumulation of epidermal cells in the 
region of the propodeum results in the formation of an especi- 
ally thick chitin layer there. Indeed, so active is the process 
of chitin secretion in this neighbourhood, that sections actually 
show minute liquid globules issuing from the chitin-secreting 
cells. 


353 


The cells, on the other hand, which are in the region of 
the future joint membranes, do not form a hard chitin, but 
produce a tough, but flexible, somewhat corrugated mem- 
brane. This is especially clearly seen in the neck region, and 
at the points of junction of the legs.with the thorax (fig. 41). 
The somatopleural mesoderm, so far as I can observe, 
always undergoes a renovation during the metamorphosis, 
and eventually persists as a delicate membrane with prominent 
nuclei, immediately below the chitinised ectodermal cells. 

The metamorphosis of the general body integument, then, 
closely resembles that described by Pérez in Calliphora. In 
that insect, however, the imaginal ectoderm extends over the 
cells of the larval ectoderm, which do not disappear till much 
later. Though cytoplasmic degeneration, somewhat similar 
to that of WVasoma occurs, phagocytosis is much more pro- 
minent, and the phagocytes attacking the integument cells 
are, generally, already strongly gorged with phagocytised 
muscle tissue (sarcolytes). 

The Phragmas.—During the pupal period the integument 
undergoes a number of changes which result in the formation 
of the phragmas—ingrowths of the integument serving for 
the inserticn of the muscles. The phragmas are of two kinds: 
there are the true phragnias, which are actual invaginations 
of the integument (fig. 43); a second type of structure which 
may be designated a “false phragma’’ is essentially an 
ingrowth of the edge of a segment into the body cavity, below 
another segment which now overlaps it. An excellent example 
of such a “false phragma’’ is the anterior part of the meso- 


- thoracic tergum, which, as already mentioned, is simply a 


prolongation of the mesothorax beneath the prothorax. The 
great phragmas of the ovipositor also belongs to this class. 
A false phragma, then, is a downgrowth of integument, which 
consists of only a single layer of cells. The true phragmas, 
on the other hand, are invaginations of the integument, 
generally hollow at first, and consist, of course, of a double 
layer of integument. They are found early in pupal life, and 
after some thirty-six hours chitinise, the chitin being 
secreted between the two layers. 

In the abdomen of the female a number of these phragmas 
are developed in connection with the ovipositor. They are 
rather short, and on them originate the great muscles of the 
ovipositor. 

The thorax and propodeum are provided with a number 
of transverse phragmas, for the insertion of the great thoracic 
muscles, and those of the legs and wings. One such phragma 
runs transversely just behind the scutum of the mesothorax ; 
below the metathorax runs a transverse horizontal phragma. 


354 


In the neck there is also a phragma, being a hoop-like forward 
and backward extension of the prothoracic shield, all round 
the neck. 

But the most remarkable of all are the great cephalic 
phragmas, which give attachment to the muscles of the 
antennae and mouth appendages (fig. 43). Early in pupal 
life a long tube-like invagination of the integument takes 
place on either side of the face, half-way between the mouth 
and the antennae. These invaginations grow upwards and 
inwards and terminate at the rear of the lower part of the 
brain. Meanwhile a second pair of tube-like invaginations 
has been formed at the rear of the head, a little below the 


_neck; growing inwards they meet the anterior pair of invag- 


inations. Since both pairs have a narrow lumen, we have the 
curious fact that at this stage a pair of long narrow canals 
run right through the head from front to rear, well above and 


either side of the mouth! The secretion of chitin, however, | 


soon takes place and the canals are obliterated, being 
gradually replaced by an exceedingly powerful rod of chitin. 

The integument on the three thoracic segments undergoes, 
during larval and pupal life, a number of remarkable changes 
which terminate in the formation of the wings and legs. The 
histogenesis of these structures will be considered first; that of 
the other appendages of the insect, which usually resemble it 
closely, can be considered more briefly. 

I'he Legs.—The imaginal discs of the legs are clearly 
visible in the earliest larvae. Here they occur, a pair in each 
of the three segments, as rather extensive but sharply defined 


areas rather thicker than the remainder of the integument, _ 


and lying on each side of the nerve cord, on the ventral side 
of the animal. The cells composing the discs are long and 
rather narrow (fig. 10). At the end of larval life the cells 
have increased greatly in number, and, as a result of the 
growth of the surrounding integumental cells the imaginal 
discs have become invaginated below the surface (figs. 2, 5, 6). 
On the base of this invagination the cells lengthen and divide 
tc form a large prominent papilla. In the resting larva this 
papilla begins to increase in size, and soon grows out of the 
invagination as a hollow appendage, dragging the mesoderm 
after it. 

Cell division by mitosis is very rapid, and the developing 
leg grows downwards, so that in the larva, before hatching, 
very well defined legs are present, hidden beneath the larval 
cuticle (fig. 3). The integument of the legs in common with 
that of the rest of the body develops a cuticle, the completion 
of which is followed by the moulting of the larva. So rapid 
has been the growth of the appendage, that its integument 


355 


becomes slightly folded in places to prevent further growth 
within the limited space afforded by the larval cuticle. This 
folding is very easily confused with the segmentation of the 
leg, a process which is not completed till some twelve hours 
later. 

The general features of the formation of the legs have 
already been described in connection with the transformation 
of the external characters of the insect. Among the most 
characteristic of these changes are to be mentioned the develop- 
rent of the bristles, spines and claws, the deepening of the 
joints, and especially the curious shrinking of the ectoderm, 
which transforms the thick ungainly appendages of the late 
larval and early pupal stages into the slender structures so 
characteristic of the imago. 

This shrinking of the legs is produced by a shortening 
and closer packing of the ectodermal cells. In the late larval 
stage these cells are long and slender, and are loosely arranged, 
but after pupation, the cells shorten considerably, become 
rather thicker, and much more closely packed together—they 
change from a loose columnar to a firm cubical epithelium. 

Segmentation of the leg in the larva, though clearly 
visible, is nevertheless little more than a series of constrictions 
due to a slight shortening of cells in this region. In the eight- 
hour pupa the constriction has become more marked, and even 
the tarsal joints are now very clearly defined (fig. 16). More 
marked segmentation is produced by a slight invagination of 
the cells of the constriction rings, a process which is rapidly 
followed by the secretion of a chitinous cuticle. In the thirty- 
six hour pupa this process has advanced considerably. 
Chitinisation continues for several days, till the whole of the 
ectodermal cells, with the exception of many of the bristle 
cells, become converted into the hard shell of the legs. 

The formation of bristles is rendered possible by the 
shrinking of the legs; it begins about eight hours after 
pupation and is practically complete thirty hours later. The 
development of bristles and spines has already been dealt 
with in connection with the development of the general body 
integument. Many of the bristles of the legs, however, take 
on a special tactile function. This is particularly clearly seen 
on the first tarsal segment of the first leg (fig. 19). On the 
lower side of this segment is a row of about twenty bristles, 
the lower ones large, the upper shorter. In suitable prepara- 
tions the lower ones can be clearly seen to be connected with 
nerve-fibres, branches of a moderately large nerve which passes 
down the leg. The bristle cell does not chitinise entirely, but 
a protoplasmic base is left, below which is a small mesodermal 
cell which attaches itself by a thin ‘‘collar” to the chitinous 


356 


integument of the leg immediately surrounding the bristle. 
From each of these cells a process is given off backwards to the 
nerve of the leg. It seems that this minute process is merely 
a neurolemma (since it stains feebly with haematoxylin, which 
the nerve fibre will not do), and this neurolemma protects 
an even more delicate nerve fibre. This nerve fibre can 
actually be seen leaving the outer part of the mesodermal cell 
and communicating with the bristle cell, within the collar 
developed from the former (fig. 19). 

I have not been able to determine exactly the function of 
the whole armature of bristles, which are so numerous on the 
legs; that they aid the insect in clinging to objects is un- 
doubtedly true, but it seems quite possible that a very large 
proportion of them have also a definite tactile function. In 
the case of the first tarsal joint this is certain; scattered 
bristles on other parts of the tarsi also have nerves connected 
with them, but the structures dealt with are so minute, that 
I cannot definitely say whether similar nerve-endings are pre- 
sent on all of them. 

Lowne has shown that the general integument of a fly 
is sensitive to touch. The same author describes bipolar 
ganglion cells lying in close contact with the tactile bristles ; 
I suspect, however, that they are really either ectodermal 
“‘receptor’’ cells, which have produced the bristle, or meso- 
dermal cells acting as a neurolemma, which encloses the delicate 
nerve fibre that terminates on the bristle. Bipolar nerve cells 
in this position are quite absent in Vasoma; but the meso- 
dermal cells which lie beneath the tactile bristles closely 
resemble such cells, the enclosed nerve fibre being very dif_i- 
cult, or often impossible to detect, partly on account of its 
. feeble staining capacity. But when the development of the 
\_ tactile organs is followed out, the nature of these cells becomes 
quite clear. In the eye they are ectodermal in origin (see 
“Organs of Vision’’); in the legs and antennae they are meso- 
dermal, and the nerves which grow towards them from the 
brain or ventral nerve cord are quite devoid of cell nuclei, 
consisting merely of nerve fibres, whose nuclei remain in the 
central nervous system. 

As Lowne did not observe the development of the tactile 
sense organs, he naturally put the more obvious interpretation 
on his observations. Indeed, it seems to be the generally 
accepted view that the cell lying below the tactile hair is a 
nerve cell; an examination of the embryology of the structures 
Epnoorneds will, however, show this view to be erroneous. I 
shall refer to this again later. 

Leucocytes, containing large amounts of débris from the 
histolysed tissues, enter the legs in an early stage in their 


357 


formation. Sometimes the narrow lumen of the leg contains 
these cells in large numbers; occasionally one may be seen 
among the loose epithelial cells of the leg. In the early pupa 
the leucocytes appear quite healthy, although heavily gorged 
with larval tissues; but gradually they disintegrate, or 
recovering, wander away, and are no longer seen here in a 
two-day pupa. The disintegration will be described more fully 
in connection with the blood. 

A single tracheal vessel enters into each leg soon after 
its formation (fig. 65). 

The formation of the muscles of the leg and of the great 
tendon will be more conveniently considered in connection 
with the muscular system. 


The development of the last tarsal segment is worthy of 
special attention. Already in the eight-hour pupa the 
segment is slightly larger and wider than those which precede 
it. As development proceeds this process continues, producing 
a claw-bearing segment considerably wider than the others. 
The ectodermal cells, moreover, do not remain as a single 
layer but proliferate, producing two masses of padding 
tissue, very similar to that which is formed in the bulging 
segment of the antennae. These masses are clearly visible in 
the pupa of thirty-six hours. The epidermal cells on either 
side of them, and also at a place on the ventral side slightly 
_ proximal to them, grow out in the form of large claws. 
Chitinisation then takes place. The cells of the three claws 
almost totally disappear; the pads, however, secrete only a 
very thin membrane of chitin, which arranges itself in two 
pairs of long pads, structures which are probably to be con- 
sidered as adhesive organs. They are shown clearly in fig. 46, 
which is from a pupa of four and a half days. The distal part 
of these pads is totally devoid of cells, the padding cells being 
confined entirely to the main portion of the segment. The 
proximal part of the padding tissue is syncitial in nature and 
on it is inserted what appears to be a tendon (fig. 46). It is 
quite possible that a pull of this tendon would draw back the 
padding tissue and apparently also the thin chitin which it 
has secreted. If the joint had been placed previously on some 
solid object it is conceivable that a partial vacuum might be 
created between the four adhesive pads and the object, thus 
enabling the wasp to cling to a smooth surface. The capacity 
of chalcid wasps for clinging to window glasses is, of course, 
well known to all who have collected them. 

The Wings.—The general features of the development of 
the wings have been described above; it remains to describe 
now the histological changes which they undergo. 


358 


The imaginal discs of the wings are seen even in the larva 
ot the first instar as two pairs of rather pronounced thickened 
areas in the second and third thoracic segments. Cellular 
proliferation takes place during larval life, and, just as in 
the legs and other appendages, the excessive growth of the 
surrounding “‘larval’’ cells produces an invagination of the 
discs; rapid mitotic cell division during the resting period 
results in an evagination of the disc, the underlying mesoderm 
being, as usual, dragged into the structure. The epithelium 
of the wing consists of a single layer of elongated cells; these 
cells, in order to present a greater surface area, frequently 
develop on their free surface in the larva shortly before pupat- — 
ing, distinct hammer-like thickenings. 

The secretion of a cuticle, simultaneously with cuticle 
development over the rest of the animal is followed by the 
pupal moult. 

The contraction of the wings, as already described, results 
simply from a closer packing of the cells. In the early pupa 
a cell of the fat-body often passes into the cavity of the wing, 
helping to nourish that structure; the mesodermal cells are 
seen undergoing mitotic division. 

After twenty-four hours the cavity of the wing has been 
almost obliterated. This is the result of at least two factors: 
firstly, the cells shorten somewhat, the peripheral pull drag- 
ging the two surfaces nearer together; secondly, they begin 
to undergo a remarkable process of wrinkling on their free 
surface, as a result of which the lower part of the cell becomes 
forced backwards. This wrinkling can already be seen com- 
mencing in the four-hour pupa; in the thirty-six hour pupa it 
is very far advanced, and the free surface of the cells which 
row present great folds, begin to secrete chitin which is itself, 
therefore, strongly folded. Many of the cells, however, in 
addition to forming folds, have also developed a hair-like 
process on their free surface; the cuticles secreted on these 
processes are the fine hairs of the insect’s wing (fig. 37). 

Other cells, again, on the anterior part of the wing 
lengthen greatly, and extending beyond the surface of the 
wing form bristles. Greatly hypertrophied cells on the hind 
‘wings produce the clinging hooks (fig 38). 

Towards the end of pupal life the greatly folded cells 
have lost the whole of their cytoplasm, this having become 
transferred, apparently, into the thin chitinous cuticle. The 
cell walls, however, are still visible, as are also the cell nuclei; 
some of these, indeed, show no visible signs of degeneration ; 
others, however, are distinctly abnormal, having lost prac- 
tically the whole of their chromatin contents (fig. 37). By 
the time the insect emerges from the pupa all the nuclei have 


359 


disappeared, except.a number on the outer edge of the wing ; 
these persist throughout the life of the wasp in a half disin- 
tegrated state; their presence can easily be revealed by stain- 
ing the wing ‘of a properly preserved insect (fig. 38). The 
outlines of the cell of the pupal wing are also clearly visible 
around the border of that of the imago; they are beautifully 
seen in the great fastening hooks of the hind wings, as long 
projections into the wing, and evidently give special strength 
to these structures (fig. 38). 

By this extensive folding of the free surface of the cells, 
the great extensions in the size of the wing take place; so. 
pronounced indeed is this, that, as already menticned, th® 
wing directly after the imaginal moult, expands to an area 
sixteen times that of the pupal wing. 

The obliteration of the cavity of the wing, as described 
above, however, is not complete; on the contrary, the first wing 
preserves an anterior (marginal) and a median longitudinal 
“sinus,’’ in the form of two great channels passing down the 
wing (fig. 44). The anterior one is bifurcate distally. The 
hind wing presents only one such channel. These channels 
are the “clear spaces’’ described above as visible in a surface 
view of the wing, and into these channels pass the tracheoles 
of the wing; leucocytes are also seen here during early pupal 
life; they disintegrate later (just as they do in the other 
appendages), but some may be seen even into the fourth day 
of the pupa. 

If a newly found pupal wing be examined in sections a 
remarkable thing 1s seen. The mesodermal cells a little beyond 
the base of the wing begin to proliferate, and then extend as 
a long column of cells right down the great fissures in the wings 
(fig. 45). Nosuch structure, however, ever extends (in Vasonia) 
into the median channel of the fore wing, though this channel 
does lodge tracheoles and leucocytes; it remains indeed 
merely as a “‘pseudo-nervure,’’ while the marginal structures 
in both wings develop into true nervures. The cells of these 
columns are elongated and ‘“‘brick-like’’ in shape; the growth 
of the column is very rapid and is complete several hours 
after pupation. Late in pupal life the internal “‘lining’’ of 
the great channels begins to chitinise slightly: the chitin 1s 
pale yellow in colour, and to this the characteristic coloura- 
tion of the nervure is due. 

To what the unfolding of the wrinkled wing A the 
emergence of the wasp is due is difficult to say. While 
not attempting to discuss its cause in all insects, I may say 
that the usually accepted view, viz., that it is produced by 
the passage of air into the tracheae of the wing, must be dis- 
carded in the case of Nasonia. Here the tracheoles are very 


ss 


360 


delicate, somewhat twisted tubes, quite incapable of altering 
the shape of the wings which bear them. It seems much more 
probable that the straightening is due to the turgidity of the 
cells of the great ectodermal extension into the channels, and 
that the wings remain firm, later, as the result of the action 
of the air on substances contained in the chitin of the 
nervures. 

In connection with the small ‘‘stigmal-vein’’ of the fore- 
wing, a remarkable structure is developed, the interpretation 
of which is very difficult. On the distal part of the veins are 
developed four rounded globules (fig. 39), the distal pair 
rather smaller than the other two. They are well known to 
workers on chalcid wasps, and are frequently used in classi- 
fications. But if a stained wing is examined under a very 
high power each of these globules is seen to contain a heavily 
staining sphere (figs. 39, 40) attached to a small conical piece 
of protoplasm, the base of which is in turn attached to a long 
fibre. The fibre from each globule passes inwards, and 
becomes lost in the substance of the stigmal vein. It seems 
probable that the whole structure is one cell, of which the 
process and the conical portion represent the cytoplasm, while 
the sphere is the nucleus surrounded by a very delicate layer 
of cytoplasm. The long process, of course, immediately sug- 
gests a sensory structure; the nerve fibre being, as usual, 
almost impossible to detect, and lying within the weakly 
staining process, which acts perhaps as neurolemma. If this 
interpretation be correct, then it is not impossible that the 
structures concerned should act as speed-detecters. Increased 
speed of flight would be produced by increased rate of vibra- 
tion of the wings; this would result in a greater centrifugal 
pull on the free (spherical) part of this remarkable structure; 
or, what is more probable, it would result in a greater fre- 
quency of the striking of this body against the walls of the ~ 
globule, and it is conceivable that this would affect the nerve 
fibre which terminated in it. 

The Mouth Appendages—The general features of the 
histogenesis of the mouth appendages are so similar to those 
of the thoracic, that a brief description will suffice here. 

The imaginal rudiments of these structures are clearly 
visible in the first larval instar (fig. 47); in their condition of 
development.the labrum and mandibles are already much in 
advance of the condition in which we find the thoracic append- 
ages; 7.¢., they are no longer merely ectodermal thickenings, 
consisting of embryonic cells, but have now become invag- 
inated well into the cavity of the head. 

The imaginal discs of the labrum are a pair of solid ecto- 
dermal ingrowths, situated at either. angle on the fore-part of 


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- —— Be. 


P ek ee. st po ae an ae Gare. be “F. a 


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- - ; “ — —~ 
sl a ae 


361 


the mouth. Their development during the larval and early 
pupal periods is quite similar to that of the thoracic append- 
ages, 2.€., the cells proliferate, grow outwards, forming a 
cavity behind them -as they do so, and drag the underlying 
mesoderm after them. The general features of their develop- 
ment have already been described. 

The mandibular imaginal discs (fig. 47) are particularly 


interesting. Each consists of a sac of ectodermal cells (lined, 


of course, with mesoderm), and invaginated well into the 
cavity of the head. The floor of the ‘“‘sac’ is many cells 
thick, the cells themselves being rather smaller than the larval 
integumentary cells. On the ‘“‘floor’’ of the invagination is 
a small number (about nine) of very remarkable cells; they 
are club-like in shape, and each bears a long cytoplasmic 
extension outwards. These processes are so arranged that 
they possess, together, the shape of the larval jaw, and it is 
from the termination of these remarkable cells that the 
minute-stylet-like mandible of the larva is secreted. This is 
seen clearly in fig. 47, which is taken from a larva shortly 


before entering the second instar. The functional jaw is no 


longer in communication with the cells which have secreted 


it; these cells, on the contrary, are now secreting a second 
mandible, within the first, and the latter will be cast off at 
the larval moult. 


We see, then, that the larval mandible is formed from the 
same set of cells which produce the mandible of the imago. 
The jaw of the larva must therefore be regarded as homo- 
logous, in part, with the jaw of the mature insect. It is, of 
course, quite conceivable that this might not have been the 
case; however, the fact that it is so can lead to important con- 
clusions, which will be considered in the second part of this 
paper. The same thing will be seen later, in connection with 
the antennae. 


During the feeding period of the larva the mandibular 


_ imaginal disc grows in size; at about the middle of this period 


the disc has become partly everted, and the projecting portion 
has the shape of the jaw of the last instar. On account of 
the much greater size of the jaw at this stage, it 1s necessary 
for a much larger part than hitherto of the imaginal disc to 


_ take part in its formation, and this is the reason for its pre- 


cocious evagination. 
During the resting stage the mandibular disc grows 
rapidly, the cells dividing mitotically. So far as I could 


_ observe, the long club-like cells of the disc in its first’ instar 


become modified during larval life into cells which do not 


_ differ visibly from the others of the madibular disc, 7.e., cells 


362 


which have become specialized into secreting a certain struc- 
ture may apparently (perhaps as a result of the act of secre- 
tion) become modified so as to resemble neighouring cells, and 
then co-operate with these in the secretion of another (some- 
times unlike) structure. The observation, if correct, would 
be of considerable theoretical importance. ‘I cannot, however, 
state with certainty whether these earliest specialized cells 
_ persist throughout larval life. 

Early in the resting period the mandibular palp already - 
mentioned above is distinctly seen. The disc grows rapidly 
by mitotic division of the cells, and drags the mesoderm 
after it. 

The remainder of the development of this, and of the 
other appendages of the mouth, closely resembles that of the 
legs, and need not be described further here. It is only 
necessary to add that the imaginal discs of the first and second 
maxillae are present in the earliest larvae, and do not differ, 
except in position (being closely applied to the mouth) from 
the leg discs. 

The Ovipositor.—The early stages in the formation of the 
ovipositor are identified with those of the legs, 7.e., the 
imaginal rudiments, present in the early larvae as ectodermal 
thickenings, become invaginated into the abdominal cavity 
(fig. 2). In the resting period they grow outwards, dragging 
the mesoderm after them; rapid cell division, by mitosis, 
results in the great extension of these appendages, along the 
ventral body wall, as has already been described fully above. 
The ectoderm of these appendages is only one cell-layer in 
thickness, and the cells themselves are generally long and 
narrow. The third appendages differ from the others, how- 
ever, in being several cell-layers in thickness; they are not 
hollow, as are the others, and are fused with the body wall. 
Into the hollow appendages, as usual, migrate leucocytes, 
which disintegrate there during the pupal stage. The second 
appendages in the larva seven hours after defaecation have 
already become closely approximated, and the cells on their 
adjacent halves have forsaken their long columnar shape and 
are now cubical; this portion of the appendage is in process of 
invagination into the outer part. 

In the early pupa, the cells of the developing ovipositor 
begin to lose their columnar shape, the characteristic of the 
growing stage, and now become cubical, and rather small in 
volume, 7.e., the ovipositor as a whole, having reached its 
condition of maximum growth, now begins to differentiate. 
The cells, themselves, become more closely packed together, 
and the long appendages shrink, just as we saw, above, in 
connection with the legs and wings. 


363 


The second appendages meanwhile have fused, and now 
enclose a cavity. Since the inner half of each of the second 
appendages has become invaginated into the outer half, it 
follows that the tube formed by their fusion must be lined 
internally by the ectoderm. 

Shrinking of the appendages continues; in the thirty-six 
hour pupa the second appendage is no longer recognizable as 
a compound structure ; it appears simply as a tube, lined by a 
single layer of ectodermal cells, the cavity containing 
mesoderm. 

During the third day chitinisation commences. The outer 
portions of the first appendages chitinise strongly; their inner 
parts, however, remain as flexible membranes, similar to those 
of the leg joints, neck joints, etc. (fig. 42). The compound 
second appendage chitinises in a very remarkable manner. 
The cells at its tip have previously arranged themselves so as 
to present a serrated tip to the ovipositor; chitinisation of this 
results in the characteristic sawing extremity. The remainder 
of the compound appendage becomes semilunar in section; the 
two outer thirds chitinise heavily, and the chitinous prisms 
so formed are connected by the median portion, whose walls 
develop into tough membranes, and enclose a quantity of 
mesodermal tissue. In side view the chitinised ovipositor 
shows a very pronounced ‘‘spiral’’ pattern, not unlike that 
of large tracheae. 

The hard chitinous sheaths and the tough membranes of 
the first and second appendages thus enclose between them a 
firm, yet pliable passage, down which the eggs pass during 
oviposition. 

An important part of the female egg-laying apparatus 
is the short appendages of the last abdominal segments; the 
general nature of these appendages has been described in con- 
nection with the general features of the insect; it remains to 
point out the nature of the tactile organs with which the 
appendage is so well supplied. 

The general features of the development of the appendage 
are identical with those of the body integument. The bristle- 
secreting cells, however, do not entirely chitinise; on the 
contrary, they seem to grow in size, and growing backwards 
from the bristle, pass well into the cavity of the appendage, 
remaining in connection with the cell only by a long delicate 
process (cf. fig. 50). A large nerve enters each appendage, 
then breaks up into nerve fibres, which communicate each with 
a bristle cell. I could find no trace of an intervening meso- 
dermal cell, similar to that described in the tactile bristles of 
the leg or antenna. The whole appendage, however, is filled 


364 


with a mass of mesodermal padding tissue, which evidently 
acts as a sufficient protection for the delicate nerve fibres. 

It seems scarcely necessary to point out again that the 
large cell lying beneath the tactile bristle is not a nerve cell, 
as 1t is usually believed to be, but that it is merely the ecto- 
dermal cell (receptor cell), from which the bristle has been 
formed. The cell is not an element alien to the bristle, but 
rather is the bristle to be regarded as a special part of the 
cell which acts as an intermediary between the cell and the 
environment, much as do the taste-hairlets at the free ends of 
the cells of mammalian taste-buds. 

The Male Copulatory Organs.—The visible cellular 
changes which underlie the formation of the penis are very 
simple; first the cells, in the early pupa, adopt the usual 
columnar (growing) shape; in the pupa two days old they 
become cubical, and chitinise a day later. 

The cells of the anterior dilation of the cavity of the 
penis, the vesicula seminalis, are quite different in shape; 
they are long and narrow, and form a thick wall around the 
vesicle. 

The Antennae.—At the front of the head are formed a 
pair of appendages, the antennae, which are quite different 
in nature from those appendages above described. Below 
them an outgrowth of nerve fibres is formed from the brain, 
on each side; and the fibres growing into the developing 
antennae terminate on modified ectodermal cells in these, — 
the modifications of the ectodermal cells into sense cells, and 
indeed, of the whole ectoderm into an antenna being of such 
a nature as to form what must be a very efficient sensory 
structure. 

The antennae are present in the earliest larvae—indeed, 
at this stage they are already more advanced than are the 
legs or wings, each being now in the form of a small papilla, 
composed of long narrow cells undergoing evagination from a 
previously invaginated antennal disc (fig. 47). They are, 
indeed, at a stage of development which the legs do not reach 
till the end of larval life. 

During the larval period the cells of the rudimentary 
disc continue to divide, so that, shortly before the larva 
defaecates, a distinct antenna is visible on the surface of the 
larva (fig. 3). Already at this stage the curiously jointed 
condition of the mature structure is clearly indicated (fig. 36), 
for the ectodermal cells have not divided regularly, as is the 
case in the legs and wings, but at short intervals a few cells 
have ceased to divide for a time, and remain long and 
columnar, while between these the ectoderm has undergone a 
marked proliferation to form a solid downgrowth of very 


we 


iz 
hued 
EA 


365 


minute cells; from the latter the tissue which produces the 
bulging of the joints is formed—a kind of padding tissue— 
whilst the former, the long columnar cells, give rise eventually 
to the joints and constrictions between the segments of the 
antenna. 

Already at this early stage a distinct outgrowth of nerve 
fibres from the brain is seen, though it does not as yet extend 
right into the developing antenna; a large tracheal vessel 
can also be distinctly seen, at this stage, within the lumen 
of the antennal projection (fig. 36). Leucocytes have also 
begun to enter. 

The mesoderm still adhering to the overlying ectodermal 
cells, follows them as they grow outward to form the antenna. 
The cells do not appear, at this stage, to have undergone 
any marked proliferation, and consequently appear as a deli- 
cate network lining the lumen of this structure. But that the 
mesoderm does eventually proliferate seems quite clear. This 
will be referred to Iater. 

The post-defaecation period of the larva is marked by a 
continuation of this process; the antenna growing rapidly 
eventually attains the size that we see in the mature insect. 
The differentiation is, as yet, however, very incomplete. The 
cells of the “‘padding tissue’’ have proliferated so rapidly that 
the mass develops a temporary invagination cavity (fig. 36). 
The columnar ‘“‘partition cells’? have divided, and now form 
a narrow ring of short cells between the masses. The ecto- 
dermal cells then partake in the process of cuticle secretion 
which is going on everywhere in the integument at this stage, 
a process which, when it is complete, is immediately followed 
by a shedding of the larval cuticle. 

The antenna now contains two tracheal vessels (fig. 65) ; 
large numbers of leucocytes are present, and the nerve out- 
growth from the brain has extended practically to the tip 
of the structure. 

A rapid differentiation now takes place. The cells com- 
prising the integument of the antenna adopt a more regular 
arrangement; the cells which give rise to the joints between 
the segments and to the proximal and distal walls of the 
segments proliferate somewhat, and, perhaps, on account of 
the presence of a hard cuticle on their outer surface, grow 
inwards, dividing the whole antenna into the segments so 
characteristic of that stage. This condition, the commence- 
ment of which is seen in the four-hour pupa, is complete in 
the pupa thirty-six hours of age (fig. 15). 

Meanwhile the formation of bristles has been taking 
place. Already in the four-hour pupa a number of integu- 
mentary cells at the tip of the antenna have elongated and 


366 


projected beyond the general surface of the antenna. This 
process is made possible by the curious shrinking, already 
referred to, which is seen in the developing appendages of the 
early pupal period. The shrinking is probably due to a closer 
packing of the integumentary cells, which transforms the 
ungainly appendages of the early pupa into the exquisitely 
moulded structures of the imago. Twenty-four hours later 
the process of bristle formation has become completed over 
the whole antenna, and the secretion of new chitin begins. 

The integumentary cells at this stage are rather long and 
columnar, and leucocytes, in various stages of disintegration, 
may be seen lying amongst them. The leucocytes of the lumen 
of the antenna are also undergoing slow disintegration, by a 
process which will be referred to later. 

At thirty-six hours after pupation the process of chitini- 
sation has become marked; it continues for a long time, but 
differs in a rather important respect from what we see in the 
chitinisation of the general body integument. The cells do 
not undergo complete chitinisation, but remain partly as living 
cells (receptor cells), with which the antennal nerve fibres com- 
municate. Thus.only the distal portion of the bristle-forming 
cells chitinises ; the proximal portion remains below and in close 
contact with the bristle which it has secreted. Even the cells 
which form the general integument of the antenna do not 
chitinise completely. 

Meanwhile the mesoderm has been undergoing remark- 
able changes. The fibres of the network already referred to 
- have increased in length; the cells have increased considerably 
in number and also in size, and they now become so disposed 
as to occupy a position below and in close contact with the 
bristle secreting cells; a very delicate connection can actually 
be seen, joining the mesodermal cell to the bristle cell above it; 
so intimate, indeed, is the communication between the two 
that it gives the appearance of a large binucleate cell (fig. 49). 


Meanwhile the nerve of the antenna: has grown in size, 
and extended as a great ‘‘tendon-like’’ axis right through the 
antenna. Covering it is a thin layer of cells, the splanchno- 
pleure of the brain. 

In each segment of the antenna this great nerve (fig. 15) 
gives off fibres, so that at the tip of the distal segment, it is 
represented only by a loose outwardly radiating bunch of 
fibres; and in good preparations it is frequently possible to 
trace a single nerve fibre from the great antennal nerve 
through the padding tissue into one of the mesodermal cells 
which lie beneath the bristle cells; that portion of the nerve 
fibre between the antennal nerve and the nucleus of the 


367 


mesodermal cell being protected by a fibre of the mesodermal 
network already described. 

The mesodermal cells must therefore be regarded as con- 
stituting a kind of neurolemma for the nerve fibres; what the 
actual connection between the nerve and the bristle cell, a 
connection which must be situated in the delicate connecting 
piece between the two cells, is, | am unable to say. These 
remarkable structures are shown in figs. 49 and 51. Very 
often two mesodermal cells lie in connection with a bristle 
cell (figs. 51, 53). The apparently erroneous interpretation 
which B. T. Lowne placed on similar cells in Calliphora has 
already been discussed in connection with the description of 
the tactile organs on the legs. 

At the tip of the antenna the bristle secreting cells have 
frequently retreated a considerable distance from the respec- 
tive bristles, and only a long protoplasmic filament remains 
to connect them (fig. 50). 

It seems scarcely possible to doubt that the structures 
here seen are tactile im nature. 

The antenna, however, is the seat of a number of other 
remarkable sensory structures. 


In the second segment, which is rather longer than those 
which follow it, there is a number of curious structures, 
which must, I think, be regarded as olfactory organs. Of 
these there are ten, and each is in the form of a long tubular 
sac, formed by five elongated cells, each with a large nucleus, 
the ten olfactory sacs hanging in a ring around the antennal 
nerve, from the distal end of the segment, into its spacious 
cavity (fig. 52). The lumen of these tube-like sacs is very 
slender, and appears to be quite devoid of chitin. It com- 
municates by a short duct with the exterior, the small circular 
openings lying in a rather deep ring-like depression immedi- 
ately surrounding the joint between the second and third 
segments. The masses of padding tissue act as supports for 
the olfactory sacs. I could not observe the innervation of 
these organs, a fact which is partly due to the minuteness of 
the nerves, and to the difficulty of staining them. Nor was 
I able to follow their whole development—in newly formed 
pupae they do not yet occur; in pupae at the fifty-six hour 
stage they have already been formed, appearing then as long 
protoplasmic sacs hanging down from the distal end of the 
segment, rather gelatinous in consistency, and not so delicate 
and slender as in the adult condition. They probably begin 
to develop, then, at about the twelve- to twenty-four hour 
stage, and there can be little doubt that they are produced 
simply as invaginations of the developing ectoderm, accom- 
panied by a great elongation of the cells concerned. 


368 


A third series of structures, which are perhaps to be 
regarded as sense cells that serve the insect in maintaining 
its equilibrium, is found in all the antennal joints, with the 
exception of the first and last. On the proximal and distal 
surfaces of the antennal segments the epidermal cells do not 
chitinise, as they do on the general body surface, but, after 
secreting a thick chitinous sheath, remain below and in close 
contact with this, as large fleshy cells, which are especially 
prominent in the angle between the lateral walls and the 
distal surface (fig. 51). Each of these large cells has the 
appearance of. being binucleate; one of these nuclei is pro- 
bably that of a mesodermal cell, the close adherence of the 
ectodermal and the underlying mesodermal cell having already 
been mentioned in connection with the cells of the tactile 
bristles. In fortunate preparations a nerve may be seen 
running to these large cells. I hope to discuss their function 
more fully in a later paper. 


A fourth series of structures, which are perhaps to be 
regarded as auditory organs, can be very clearly seen on the 
last nine antennal joints. They are confined to the female, 
and each structure is‘ in the form of a long, rather narrow, 
hollow cylinder, sharply pointed distally, and formed of thin, 
clear, transparent chitin (fig. 53). Immediately beneath 
these hollow cylinders lies a mass of fleshy cells; sometimes 
as many as five nuclei are visible, and no distinct cell walls 
can be recognized. What is apparently a strand of nerve 
fibres can occasionally be seen entering this cell mass. 


These organs are very prominent on the antenna of the 
female, each being nearly as long as the antennal segment 
bearing it, and frequently projecting in a sharp point beyond 
it. The number in the several segments varies; the first and 
second bear six each; the third has eight; the next three, ten ; 
the seventh, twelve; the eighth, fourteen; and the last, eight. 


Only the first (proximal) segment of the antenna is pro- 
vided with muscles (fig. 11). These run longitudinally; dis- 
tally they are inserted into the upper portion of the tip of 
the first joint; then passing backwards, they diverge a little 
and, entering the head, pass downwards, and become inserted 
on the great cephalic phragmas. The lower portion of the 
wall bounding the opening in the head through which the 
nerves, muscles, and tracheae pass into the antenna serves as 
a pulley on which the antennal muscles work. 


A second set of muscles is confined to the antenna; it has 
its origin along the posterior ventral half of the first joint 
and is inserted into the upper part of the base of the second. 
The function of this system is, obviously, to raise the greater 


369 


part of the antenna, irrespective of the action of the other 
set of muscles. 

The muscles begin to develop in the pupa of four hours; 
the actual histogenesis of these muscles, which does not differ — 
from that of other muscles, will be described later. 


By this process, then, there is formed the antenna of 

the imago, a structure which is to be regarded simply as a 

highly sensitive portion of the integument, modified and 

: grown out in such a way as to permit of a maximum of 
_ efficiency in the action of the sense cells, which it bears. 


’ 


The Organs of Vision. 


The Compound Eye.—But of all the changes undergone 
by the ectoderm as it gradually develops in the larva and 
| the pupa, the most remarkable are those which take place at 
| the sides, and in front, of the head. Here the ectodermal 
cells become exceedingly specialized, and, while retaining 
their primitive function of acting as a protection for the 
internal organs, as well as, to a certain extent, their capacity 
for secreting a cuticle, yet become modified, and disposed 
in such a manner that the terminations of outgrowing nerve 
fibres from the brain, which come to end in close relation with 
them, may become stimulated in certain ways by the light 
rays emitted from external objects, the vague impressions of 
which become modified, as a result of their physical media- 
tion, into what must now be very highly specialized sensations. 


; 
| 
| 
As a result of the processes, which begin in the embryo, 
| and are continued right throughout larval and pupal life, the 
great compound eyes and the three smaller ocelli become 
developed. 
The formation of the compound eyes will be considered 
| first. Already in the larva of the first instar the ectoderm 
of the head, on either side of the brain, has begun its modi- 
fied course of development. The ectoderm at this stage con- 
sists of a large number of cells disposed roughly in three layers 
(fig. 55). Although no examination of eyes in unhatched 
embryos was made, yet there can be no doubt that the cells 
of these layers are formed as a result of a division of vertically 
elongated cells, the disposition of the cells at this stage being 
such as to indicate that they had arisen from those of the 
middle layer. Several individual cells are shown isolated in 
fig. 54 for greater clearness. The cells of these three layers 
can already be distinguished morphologically. Those of the 
external layer are rather short and generally conical in shape; 
the middle layer cells are long and generally spindle-shaped, 
with the nucleus in their middle, while the cells of the inner 


=. 


—— a a n 
. 


370 


layer are elongated, broad and conical at their bases, and 
prolonged externally into a long rather narrow process; the 
nucleus is confined to the lower conical portion of the cell. 
In all three types of cell the nuclei are alike; there is a fairly 
distinct nuclear membrane, and the chromatin is contained 
in a sharply defined karyosome. The cell cytoplasm is devoid 
of granules. 

During larval life there is a great proliferation of the 
cells of the imaginal disc, unaccompanied, however, by any 
marked visible differentiation; so that the optic disc in the 
larva at the time it ceases to feed is little different from the 
structure as we see it in the first instar, except that thé cells 
are ever so much more numerous, and actually smaller than 
in the early larva. JI could find no evidence of renewed 
differentation of optic disc cells from unmodified head ecto- 
derm during larval life, such as occurs, according to Giinther 
(1912), in the developing eye of Dytiscus marginalis. On 
account of the great crowding together of cells at this stage, 
it is very difficult, except in places where they have been 
accidentally loosened, to observe the actual structure of the 
individual cells. No marked difference can, however, be 
noticed between these cells, and those of the early imaginal 
disc. 

About the time when the larva ceases to feed, the cells 
begin, as a result, probably, of their mode of division, to 
adopt the arrangement in groups somewhat as we see them in 
the adult wasp. The basal cells, with a very elongate oval 
nucleus whose chromatin is arranged in scattered granules, 
which apparently develop the rhabdome, can now be seen 
extending right to the external surface, and the cells of the 
middle layer are seen to surround this cell in groups of seven. 
These are the sheath cells which, with the basal cells, form 
the developing ommatidia. At the time when the larva begins 
to defaecate the cells of the external layer have extended 
throughout the thickness of the disc, and can be seen under- 
going longitudinal (vertical) fission, their nuclei being retained 
in their outer portions (fig. 56). At their bases (distal ends) 
can be seen, in good preparations, four: minute cells, which 
have probably been budded off from them. These are the 
undifferentiated vitreous cells, which later become so prom- 
inent. In the larva at the time of defaecation, a single pair 
only, as a rule, of the elongated outer-layer cells can be seen 
between adjacent ommatidia. Their disposition is such as to 
show very clearly that they have quite recently undergone 


(1) They are, as a rule, spoken of as “‘retinula cells.’’ This is, 
however, due to a misconception of their function, and they will 
here be spoken of simply as ‘‘sheath cells.” 


ee 


371 


longitudinal fission (fig. 56). At times, though very rarely, 
a third such cell can already be seen connected with the devel- 
oping ommatidium. The most obvious feature of the develop- 
ing compound eye at this stage, and for the whole of the 
next day, is the closeness with which the cells are disposed, 
making accurate observation of the development impossible 
except in places where the cells have become artificially 
loosened. 


This process of longitudinal division of the cells surround- 
ing the ommatidia continues for a time after defaecation, till 
four such cells are formed round each. In larvae eight hours 
after they have defaecated this process is complete (fig. 57). 
At this time there have also been formed, almost certainly 
from these same outer-layer cells, a pair of rather small clear 
cells, developed at the outer end of each ommatidium, and 
generally very distinctly visible; they do not attach them- 
selves as closely to the ommatidial cells as do the others. 
The four long cells which embrace the ommatidia are the 
developing pigment cells; during their formation from the 
outer-layer cells their nuclei have taken up a more internal 
position. The two small cells lying external to them will 
become the lens cells. 


At this stage, then, the optic disc consists of a great 
number of developing ommatidia, each consisting of a large 
central basal cell, closely surrounded by seven sheath cells, 
while at the distal end of each there are four vitreous cells, 
and two lens cells outside these, adjacent ommatidia being 
separated by the four elongated pigment cells which-surround 
these structures. 


The processes described so far have consisted almost 
entirely in the cells adopting the position in which they 
occur in the adult; visible differentiation has not proceeded. 
beyond the rough assumption of size of the adult cells. The 
remainder of the development consists of a change of the 
general disposition of the optic disc as a whole (due chiefly. 
to an increase in the length of the cells), and of a partial 
disappearance and gradual transformation of these almost 
undifferentiated cells to the condition in which we find them 
in the adult. 


The former process will be considered first. In the larva 
in its first instar the imaginal disc of the compound eye is 
very prominent, forming a definite thick area at each side of 
the brain; as the larva gradually develops, the cells, as we 
have seen, divide very extensively; hence the disc becomes 
larger in area, but the cells, not having increased in size, are in 
no way any more distinct ; indeed, as the ectodermal cells 


’ 


372 


surrounding the developing eye have been increasing in length 
during this process, the disc is actually less distinct than in 
the larva in its first instar (fig. 78). So marked has been the 
disparity in growth between these two parts of the ectoderm 
that in the large larva, before the optic disc cells begin to 
grow in size, the disc has undergone a distinct invagination 
by the partial growth over it of the unmodified head ectoderm. 
From now on, however, the disc gradually thickens, while the 
ordinary ectoderm of the head becomes rather thinner. In 
fig. 77, which is from a larva about sixteen hours before 
pupation, we. see the disc already sharply marked off from 
the rest of the ectoderm. In fig. 79, which is taken from a 
pupa thirty-six hours of age, the cells have elongated greatly, 
and are beginning to turn inwards, towards the optic nerve, 
as it grows out from the brain. Fig. 80 shows a section of 
the eye of a pupa which is about ready to emerge. The cells 
have increased greatly in length, and the bases of the 
ommatidia converge upon the optic nerve. Meanwhile there 
has been a gradual increase in the convexity of the eye. (In 
fig. 78, which is taken from an advanced larva, the eye is 
shown as very much more convex than in the next stage, 
taken some ten hours later. There is, however, no real com- 
parison between the two, since the first is from a still actively 
moving larva, in which the flexible disc must necessarily 
be subject to considerable distortion.) 

The change in direction of the ommatidia probably finds 
its explanation in the following observation. Beneath the 
optic disc lies a series of tissues: the mesoderm of the body 
wall (somatopleure), a membranous ingrowth of ectoderm, as 
will be described later, and an outgrowth from the brain. @) 
These three form a fairly thick mass beneath the optic disc, 
which later becomes exceedingly firm by the deposition of 
chitin. Now, as the cells of the eye gradually increase in 
length they will necessarily be subjected to pressure from 
these membranes below, and from the cuticle, which they have 
secreted, above. The cells of the eye, under these circum- 
stances, will be able to retain their straightness for the greater 
part of their length, which is absolutely essential for them, 
only by growing in the direction of least pressure, 7.e., towards 
the middle of the disc; hence whereas the cells in the middle 
can remain vertical, those which are some distance from the 
middle will have to take up a more oblique position, while 
those at the circumference of the optic disc, where the cuticle 
and somatopleure virtually adhere, will necessarily have to 
take up a horizontal position if they are to develop at all; 
in short, the degree to which the cells converge will depend 


(2)The nature of these membranes will be discussed later. 


373 


upon their distance from the central ommatidium. Under this 
pressure, of course, not only the somatopleure and its adjacent 
membranes, but also the cuticle, will bend, and this would 
account for the increase of convexity of the eye as the cells 
gradually grew. (The obvious question as to why, under this 
pressure, the cells might not be expected to converge just as 
readily towards the exterior as in the opposite direction finds 
its reply in the fact that the ommatidia are much broader 
at their distal than their proximal ends; indeed, they are 
really cone-shaped structures, so that such an arrangement 
would not be possible.) 

A short digression may be of interest here. If the above 
suggestion is correct, it will follow that the convexity of the 
eye of Vasoma depends upon the ratio between the tension 
of the somatopleure and its adjacent structures and that of 


the cuticle covering the eye. Insect eyes vary greatly in 


convexity; one has but to compare the almost spherical eye 
of a Cicada with the rather flat eye of many flies. If it 
should be of advantage to a species to have an eye of greater 
or lesser convexity, it follows that it would not be necessary 
to postulate, in the germ cells, a factor for increased eye 
convexity, but that the result could be obtained simply either 
by a strengthening of the germinal representative of the 
membranes underlying the optic discs, or by a weakening of 
that of the optic cuticle. 

It is necessary now further to consider the histological 
changes undergone by the developing eye. In transverse 
sections of the ommatidia the rhabdome cell is seen to be 
fairly thick, especially at its base, where its nucleus lies. 
The seven sheath cells can generally be clearly made out, 
surrounding it (fig. 58c). The pigment cells are long, and 
extend through the thickness of the disc; the nucleus is in the 
middle of the cell, although the distal end is still generally 
the widest part of it. The four vitreous cells have now 
become fairly distinct; occasionally the lens cells appear to 
be differentiating, presenting at times a rather vacuolated 
appearance. They are also seen embracing the outer end of 
the visual cells more closely. 

But it is not till after pupation that any really marked 
changes appear. In the pupa of about four hours the cells 
of the developing eye have already increased considerably in 
length, the thickness of the disc being now 30p, of which 
about 254 represent the length of the sheath cells. The 
rhabdome cell has meantime narrowed considerably, the 
proximal end, in fact, having become developed into a rather 


_long narrow filament. The nucleus, which is situated in its 


lowest portion, is visible only with difficulty. Distally it 


374 


extends right to the outer surface, where it sometimes pro- 
jects as a distinct button-like structure. In longitudinal 
sections of the ommatidium the outlines of the sheath cells 
are now very difficult, or almost impossible to detect. But 
if such ommatidia are observed in sections cut transversely 
to their length, the central rhabdome cell and the seven sheath 
cells surrounding it can usually be clearly seen (fig. 58d). If 
the ommatidia are examined in pupae a little older, we find 
that the number of sheath cells has been reduced to six 
(fig. 60). Grenacher, working with Dytiscus marginalis, and 
Johansen, using Vanessa urticae, could find only six sheath 
cells. On the other hand, Hesse regarded seven as the normal 
number for Arthropods; Kirchhoffer found this number in 
Dermestes vulpinus. Giinther (1912), using the same material 
as Grenacher had employed much earlier, found seven sheath 
cells in Dytiscus marginalis. Seven is then, evidently, the 
number of sheath cells occurring in the early stages of the 
insect eye. 
According to Giinther, this number is reduced to six by 
one of the cells becoming pressed out from among the others. 
~_I believe the same thing occurs in Vasona in the early pupal 
| | period, but the structures dealt with are so exceedingly minute 


in this insect, that accurate observations on this point are 

_very difficult. I am also unable to describe the ultimate fate 
of the seventh cell that has been cast out, whether it dis- 
integrates, or whether the other sheath cells develop at its 
expense, or, finally, whether leucocytes absorb it. 

The four vitreous cells are rather distinct, and the devel- 
oping lens cells are continuing to apply themselves more 
closely to the distal end of the ommatidium. The six pig- 
ment cells have become very narrow, their nuclei remaining 
in a position considerably above their middle; at times the 
pigment cells have still a distinctly spindle-like appearance. 

The visible changes that take place during the next 
twelve hours are not very pronounced; the filament-like pro- 
cess at the proximal end of the rhabdome cell becomes more 
marked; at the periphery of the optic disc it undergoes an 
extraordinary elongation, becoming about two-thirds as long 
as the rest of the ommaditium (cf. fig. 79). The distal end of 
the cell projects quite distinctly beyond the vitreous and 
lens cells (fig. 61); 1t is possible, however, that the appearance 
of this structure in preparations is due to the action of re- 
agents used in making them. The vitreous cells have become 
quite distinct, and the two lens cells have embraced them still 
more closely. Their protoplasm has become slightly granular, 
while the nucleus is very large, with scattered chromatin and 
a very distinct though small nucleolus, and lies in the lower 


; % hi] 
Py 
AA 4 
ai 

/ 


375 


part of the cell. The four pigment cells) surrounding the 
ommatidium have altered in shape; they appear now as rather 
thin filaments, with a great swelling at a distance of about 
one-third their length from the anterior end, a swelling which 
lodges the nucleus (fig. 59). The latter is rather distinct; its 
chromatin is scattered, and it contains a small but very dis- 
tinct nucleolus. 

From now onwards the visible changes become more pro- 
nounced. The rhabdome and sheath cells continue to grow 
in length, reaching in the twenty-one hour pupa a length of 


- about 38y, the total thickness of the eye at this stage being 


it 


48u. The four vitreous cells have increased in size, and now 
quite surround the end of the rhabdome cell except at its 
termination, where it can still often be seen projecting beyond 
them (fig. 61). Their cytoplasm is fairly clear, and they 
possess each a relatively very large, rather irregular nucleus. 
The two lens cells, which have been gradually approaching 
the vitreous cells from the sides, now wholly surround them 
above and at the sides, embracing them closely (fig. 62). The 
nuclei are lodged in their lower portion; each possesses a 
very distinct nucleolus. The rhabdome cell has meanwhile 
been narrowing for the greater part of its length, but it is 
even yet visible, though only very faintly, amongst the 
vitreous cells. At this stage the sheath cells which have 
extended by very delicate processes, over the proximal filament 
of the rhabdome cell, begin to develop granules of a reddish- 
brown pigment throughout their length, so that in this insect 
the capacity of forming pigment is not confined to the true 
pigment cells (fig. 63). The latter, indeed, at this stage are 
still quite devoid of granules. Other changes, however, which 
may be the forerunners of pigment formation, are now going 
on in them, and they now assume a very remarkable form; 
the cells which were, before this, filamentous, or at times 
spindle-shaped, become even narrower, except in their distal 
third, which remains rather thick (fig. 62); their protoplasm 
becomes vacuolated, the vacuoles at times producing small 
swellings in the thread-like filament. It is towards the 


proximal (inner) end of the cell, however, that this process 


has its most remarkable result. Here, at a short distance 
from the end, a relatively huge vacuole is formed which causes 
this part to swell up in a large globule. This condition is 
very characteristic of the pigment cells at this stage. During 


(3) At first sight, there appear to be six pigment cells sur- 
rounding each ommatidium. Each pigment cell, however, becomes 
applied to two ommatidia; hence of the six which are observed 
around each ommatidium, two ‘‘belong’’ to adjacent ommatidia. 
This will become clearer by examining fig. 64d. 


ae | 


376 


the next two days the globule remains, and, if anything, 
grows even more distinct (figs. 59, 61, 62, 73). The nuclei 
do not change their position; their nucleoli, at times, become 
relatively gigantic. 

From now onwards the cells do not increase much more 
in length, the eye reaching (exclusive of the lens) a thickness 
of about 50u in the adult wasp. The most clearly visible 
changes during this period that occur in the cells are those 
connected with the deposition of pigment in the pigment cells. 
This takes place during the third day of pupal life. The 
slender filamentous pigment cells become highly vacuolated ; 
indeed in many, at this stage, the vacuoles occupy so great a 
space and have adhered to such an extent, that practically all 
the cytoplasm of the cell lies at the periphery (fig. 73). It 
has also been seen that whereas the greater part of the pig- 
ment cell is in the form of a filament, there is a very char- 
acteristic globular swelling a short distance from its proximal 
end, while the distal third remains thick. The nucleus now 
moves upwards a little, and lodges in the distal thickening, 
so as to lie close beside the lower portion of the adjacent lens 
cell. At the same time the distal end spreads out and 
embraces portion of the lens cell nearest to it (fig. 64). The 
pigment cells thus co-operate in forming a complete coat 
round the vitreous and lens cells. The cells now begin ‘to 
undergo pigmentation. The distal fifth, on account of its 
thickness, forms a heavy mass of “‘iris-pigment,’’ evidently 
rendering the vitreous and lens cells opticaliy isolated from 
cne another (fig. 64d). This isolation is increased by heavy 
pigmentation of the lens cells in the pupa of three and a half 
days. In the proximal third of the cell pigmentation is fairly 
heavy (fig. 64b), but much less so than in the distal portion. 
The region of the pigment cell intervening between these 
two parts presents only a single row of reddish-brown granules, 
enclosed in a very delicate sheath—the cell membrane. The 
increased formation of pigment in the proximal portion of the 
cell must be connected directly with the globule which forms 
here, and which, in the late pupa, has quite disappeared. 
We thus recognize three differently pigmented layers in the 
eye of Vasonia; an outer heavily developed ‘“‘iris’’ pigment 
layer, an intermediate weakly pigmented layer, by far the 
thickest of the three, and a rather small, fairly heavily 
pigmented lower layer (fig. 80). The granules are exceed- 
ingly minute, and vary in shape from spherical to almost 
cylindrical. 

Meanwhile the sheath and rhabdome cells have continued 
their development. Transverse sections of the eye show 
numerous ommatidia cut across. Each of these is seen to 


a 
Alea 


377 


consist of the central rhabdome cell, surrounded by six sheath 
cells all embedded, it appears, in a gelatinous(?) matrix. 
In almost mature pupae the rhabdome cell appears as a 
brownish rod; workers with larger insects seem to agree that 
this consists of chitin, and the appearance of the rod in 
Nasoma, with its sharply defined outline, certainly lends sup- 
port to this view. The rod has evidently been secreted 
directly from the rhabdome cell. (This view has, of course, 
been assumed throughout in applying the name to that 
cell().) It can be seen entering the distal cell complex, but, 
as far as I could observe, does not reach to the exterior. 


The development of the lens cells has already been 
described. The development of the vitreous cells which they 
enclose is completed at about this time. This consists in a 
curious movement of their nuclei upwards to lie very close 
below the chitinous lens, and the four cells arrange them- 
selves in such a way that an opposite pair is in contact for 
a considerable. distance, so that the remaining two do not 
meet each other (fig. 64c), but so far as I could observe, the 
rhabdome no longer extends right through this distal cell 
group. 

Meanwhile the cells have not lost their property of 
secreting cuticle. During larval life the outer layer of cells 
of the optic imaginal disc, from which the pigment cells, 
vitreous cells, and lens cells later develop, secrete the larval 
cuticle (fig. 55), while towards the end of larval life they 
secrete the cuticle of the pupa. But when once ‘this cuticle 
has been secreted, the cells commence to differentiate into 
pigment cells, vitreous cells, and lens cells, and it is in the 
last alone that this property of cuticle secretion is retained. 
Already in the pupa of twenty-one hours, at a time, namely, 
when the lens cells have scarcely surrounded the vitreous cells, 


an outwardly convex cuticle is being secreted by each 


ommatidium. Since the rhabdome cell at this stage forms 
part of the external boundary of the ectoderm, it seems diff- 
cult to deny that it plays a part in this process. Indeed, 
since the rhabdome cell secretes a chitinous rhabdome over 
the greater part of its: length there is no apparent reason 
why the distal part, included among the vitreous cells, should _ 
lose this property. If we grant that this portion of the cell 
assists the lens cells in the early stages of lens formation, we 
might have a suitable explanation for the otherwise unex- 
plained disappearance of the rhabdome cell from the end of 


- (The sheath cells are usually regarded as aiding in the 
secretion of the rhabdomes.. I could, however, find no evidence 
for this in Nasonia. 

L 


378 


this cell complex. But the chief agents in the formation of 
the lens are the lens cells, which gradually transform the 
rather thin concave-convex cuticle of the early pupa into 
the thick biconvex mass as we see it in the late pupa and 
adult (fig. 64). 

By this remarkable series of changes, the originally 
three-layered condition of the imaginal disc of the first larval 
instar, which has itself doubtless been produced from a pre- 
viously single-layered ectoderm, gradually transforms itself 
into the wonderfully specialized state in which we see it in 
the adult—a state in which it does not depart essentially 
from the three-layered condition, and in which the function 
of secreting cuticle is retained, though modified, in such a 
way as to aid it in performing its new function. 

A feature of the compound eye of Vasonia is the entire 
absence of tracheae between the ommatidia, structures which 
are so prominent in the eye of Calliphora (Lowne, Hickson). 

The description of the compound eye has been confined, 
so far, to a consideration of the development and differentia- 
tion of the simple optic disc of the newly hatched larva. But 
immediately below the eye there is formed during the late 
larval and pupal life an important structure whose function 
it is to support the nervous elements of the eye. This struc- 
ture is developed directly from the ectoderm surrounding the 
optic imaginal disc, and it will be necessary to describe its 
development here; it will also be convenient, though not 
perhaps strictly logical, to give. an account at this stage of 
the development of the innervation of the compound eye, 
since these two processes are intimately connected with one 
another. 

An examination of a medium-sized larva shows that 
immediately below the ectoderm there is a delicate membrane 


' with distinct nuclear swellings, the mesodermal somatopleure ; 


it is clearly visible below the optic disc, and no other tissues 
underlie it. But when the larva is about to defaecate it is 
seen that two areas of proliferation have arisen in the head 
ectoderm above and below, and in close contact with the optic 
disc. From these areas the proliferating ectodermal cells 
grow towards each other, and finally meet, forming a very 
prominent bridge across the back of the imaginal optic disc. 
The cells soon spread out laterally, and form a membrane 
completely covering the back of the developing eye (figs. 77, 
78). Internal to this disc, of course, the somatopleure must 
lie. In the larva which is about to pupate the membrane has 
extended completely behind the eye. When the membrane 


_ 1s examined at this stage a very remarkable thing is seen. 


The cells stand off a considerable distance from the adjacent 


rr 


I — 
age 


379 


optic disc, so that a very distinct basement membrane can 
now be seen underlying it. This membrane connects the 
internal énds of adjacent pigment cells, from which it has 
doubtless been secreted. These pigment cells surround the 
ommatidia fairly closely, but are not in direct contact with 
them, and they secrete the basement membrane in such a way 
that a hole is left at the base of each ommatidium, thus per- 
mitting the easy entrance of a nerve towards the rhabdome 
cell. This basement membrane is the fenestrate membrane 
of the eye, and will be referred to as such hereafter. It 
undergoes scarcely any visible change during the rest of larval 
and pupal life. 

The cells of the inflected ectodermal layer (which may, 
for convenience, be spoken of as the periontic membrane, to 
show its relation to Hickson’s Periopticon) now undergo a 
remarkable process of branching (fig. 74), the branches being 
of three types: (1) those which unite the cells with similar 
processes from other cells of this layer, (2) those which connect 
the cells with the fenestrate membrane, and (3) those wheal 
grow in towards the brain. 

The cells themselves are large, with a distinct but irregular 
nucleus; the branches which connect neighbouring cells 
together are not very numerous. The second type of process. 
is very remarkable, and is seen to join up, each, with the 
base of a group of pigment cells, and several such processes 
may be seen coming from a single cell of the perioptic layer. 
Since these processes thus fuse in reality with the circum- 
ferences of the holes in the fenestrate membrane at the bases 
of the rhabdomes it would seem probable that they are 
hollow; later events show that this must be so. The third 
type of branching is also very remarkable; this consists 
essentially of a great massive outgrowth of fibres towards the 


adjacent cortex of the brain. This great fibrillar mass from 


the inner side of the cells of the perioptic. layer now enters 
the more ventral portion of the brain (having apparently 
broken away its own somatopleure and the splanchnopleure 
of the brain) and gradually terminates amongst the cortical 
cells comprising it. At the same time these cortical cells 
become active, and, dividing mitotically, begin to proliferate 
and to migrate outwards in the meshwork of fibrillae, towards 
the optic disc. | 
The function of the perioptic membrane is thus to form — 
a kind of-neuroglia to support the nerve cells of the optic 
ganglion; but it seems to have a second function, namely, to 
act as what must essentially be regarded as a neurolemma. 
From the above description it follows that no nerve fibre can 


penetrate to the rhabdomes unless it can enter the cavity of 
L2 


380 


the processes which are attached to the fenestrate membrane. 
I was unable to observe how the nerve fibre penetrates to 
this portion, z.e., whether 1t forces its way through the cyto- 
plasm of the cells of the perioptic layer, or whether, 
entangled as it is in the fibrillae of the cells, it works its way 
just below the cell membrane, and passing round to the 
opposite side, enters the process to the fenestrate membrane. 
It is certain, however, that a fibre from a nerve cell does 
eventually work its way into one of these processes. This is 
clearly shown in fig. 74, where the nerve cell gives off a long 
fibre which communicates with the ommatidium, and is 
entirely enveloped by the cell process which meets the fene- 
strate membrane. The cells of the perioptic membrane must, 
therefore, be regarded as functioning, also as neurolemmae. 
Moreover, it follows that, as a single cell gives off processes 
towards a number of ommatidia, a single perioptic cell must 
act as neurolemma for a number of distinct nerve fibres. I 
have not been able to see distinct instances of this in my 
preparations, partly because, in pupae a little older, when 
this process has been completed, the cells of the perioptic 
membrane have cohered closely together, making further 
observations on this point impossible; but the fact that all 
the ommatidia, several of which were supplied with processes 


from a single perioptic cell, later have nerves entering them, © 


leaves us no alternative but to accept this view (compare, 
however, fig. 75). The nerve cells which have entered the 
perioptic membrane can often be seen to give off a distinct 
process backwards. towards the brain; but I was quite unable 
to trace any of these fibres to their termination. 

The coherence of the cells of the perioptic membrane, 
which takes place in pupae a few hours later, gives the struc- 
ture a much firmer appearance, the loose branching network 
of the pupating larva being transformed into a fairly thick 
pavement membrane. At this stage leucocytes are occasion- 
ally seen between the perioptic membrane and the optic disc. 
What their function is I am not able to say. 


When pupae about twenty-four hours old are examined, a 
further development of the perioptic membrane is seen to have 
taken place. The ‘‘neurolemmal]”’ processes are no longer visible ; 
probably the best interpretation which can be placed on this 
is that their disappearance is only apparent, and that they 
have now assumed their true function as neurolemmae and 
have closely enveloped their respective nerve fibres (fig. 75). 
These fibres can be seen communicating with every omma- 
tidium, but they are so exceedingly minute that the non- 
appearance of a neurolemma as distinct from the nerve is 
only to be expected. I am also unable to say where the nerve 


ee 


381 


+ 

ends and where the ommatidium starts; and whether the 
"nerve terminates on a rhabdome or on the sheath cells. The 
fact, however, that the latter have undergone pigment 
degeneration, and the close resemblance of the sensory cells 
of the ocelli (which are innervated) to the rhabdome cell, 
: seem to indicate that it is in the rhabdome of the ommatidium 
that the nerves terminate. 

At this stage the fenestrate membrane shows a curious 

appearance which may easily be misinterpreted; the ends of 
the pigment cells which have secreted it become dilated into 
‘small cones, and give the membrane the appearance of cellular 
tissue; in reality, however, it is entirely non-cellular (cf. 
fig. 62). ; ' 
The development of the great mass of fibrillae from the 
side of the perioptic membrane towards the brain, as above 
: described, is confined to the middle third of that membrane; 
' consequently the complete optic ‘‘nerve’’ never occupies an 
area greater than this. In the thirty-six hour pupa the fibrillae 
_have become so massed together as to form a thick layer 
of fibres running longitudinally to and beneath the disc, the 
individuals of which are no longer visible. These, then, pass 
_ down the optic nerve and enter the brain. 

In the pupa of the third-day pigment granules begin to form 
in the optic “‘nerve,’’ and become very prominent a day 
later. Changes are now taking place which involve the 
ommatidia as well; these consist of a gradual alteration 
: of the shape of these structures. In the pupa at about the 
end of the third day that portion of the perioptic membrane 
“which has not been concerned in the formation of the optic 
Beerve begins to undergo chitinisation, and since this mem- 
_ brane was produced from, and still is continuous with, the 
ectoderm immediately surrounding the eye, it follows that 
_ this chitin layer will be similarly continuous with the chitin 
which now begins to form on the head of the wasp; this 
_chitinisation of the back of the eye appears to push the 

nervous part of the optic ‘‘nerve’’ out of position, by pressing 
on its periphery; at any rate, it now assumes an outwardly 
convex form. This effect is really produced by a “‘shear-like”’ 
action of the chitinising perioptic membrane, the lower por- 
tion pressing outwards and upwards, the upper down and 
“inwards. This process is complete in the four and a half-day 
' pupa (fig. 80). The ‘‘optic nerve’ from the brain at this 
b stage has also assumed the appearance of a solid projection 
from the lower side of the brain in its more ventral portion 
[ and is crowded with nerve cells; the detection, however, of 
Individual nerve fibres in this region is quite impossible owing 
_ to the close coherence of these. 


7 


382 


This shear-like action of the chitinising perioptic mem- 
brane results also in a curious change in the shape of the 
ommatidia, which is very easy to recognize (in spite of their 
very close clustering together in this region), on account of 
the rows of red pigment granules which run along them 
(pigment of the sheath cells). The lower ommatidia, which 
were originally straight, now become bent, and, as the pass- 
ing inwards and upwards of the perioptic membrane 
increases, become bent more and more, and eventually come 
to curve back upon themselves, in order to maintain con- 
nection with the optic nerve. This recurving is exceedingly 
characteristic of the lower ommatidia of the eye; from the 
above description it necessarily follows that the higher the 
ommatidia are in the eye, the less will they be bent; in the 
upper ones, indeed, the bending has been only very slight 
fig. 80). 
Ne The outstanding feature, then, of the development of the 
eye during the last day and a half of pupal life is the bending 
outwards and compression of the optic ‘‘nerve,’’ aud the con- 
sequent curving of the ommatidia, movements which are pro- 
bably to be explained as due to the compressing action of the 
perioptic membrane, as it begins to chitinise. 


By this complex process there is gradually produced the 


eye as we see it in the adult wasp, with its corneal lenses, 


pigment layers, and ommatidia resting upon the fenestrate 
membrane, which admits the fibres from the optic “‘nerve,’’ 
and in intimate relation with which has been produced a disc 
of chitin which protects the eye from within, and the whole 
organ covered internally by a very feeble membrane—the 
mesodermal somatopleure. 

It is necessary to refer now to the work of others on the 
development of the eye. It is in Weismann’s great memoir 
(1864) that we find the first correct account of the develop- 


ment in its main outline. He regarded the layer of lenses’ 


and ommatidia as arising directly from the surface ectoderm, 
while the optic ganglion (‘‘bulbus’’) he regarded as being a 
direct outgrowth from the brain. He apparently even saw 
the perioptic membrane, of which he says: ‘‘Between the 
bulbus and the disc there penetrates a thin layer of fat and 
granule cells, from which the cells which unite the two surfaces 
very probably develop.’’ He summarises his description thus: 
“The morphological value of the different parts of the eye is 
as follows: the cornea is the chitinous skeleton; the other 
parts of the eye-chamber (the crystalline cones, nerve rods, 
and their investments) are modified hypodermis; all the 
central structures (the ganglion layers and bulbus) are formed 
as outgrowths from the nervous system’’ (quote from Lowne). 


| 
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‘ 
\ 


383 


elf the perioptic membrane in Nasonia and the blow-fly are of 
similar origin, then Weismann’s view as to its origin is in- 
correct ; his very recognition of the membrane, however, in 
g hand. dissections of fly pupae is itself a remarkable instance 
: of his power of observation. His descriptions are supported 
by the work of G. H. Parker (1890) and of Carriére (1884). 
: Later workers, using much more accurate methods, have con- 
_ tradicted Weismann’s statements; their descriptions, unless 
: the process is different in the material which they used from 
what we see in Wasoma, are, however, quite erroneous. 
: A number of writers, a ee Reichenbach (1886) or Patten 
(1886), regard the Arthropod eye as having arisen as an 
_ invagination of the ectoderm, with subsequent fusion of the 
rim of the depression. The upper and lower layers of the 
invagination then meet and produce, between them, the 
vitreous and lens cells and ommatidia. According to Reichen- 
_ bach (working with the crayfish), two other layers are formed 
_ between these two. The-superficial and outer of these two 
layers then fuse and produce the layer of vitreous and lens 
cells; the third layer forms the rhabdomes, and the inner 
layer is actually regarded as forming the ganglion. 
| Patten (1886), on the other hand, regards the superficial 
_ layer as forming the cornea; the outer layer of the flattened 
_ vesicle disappears, and the rest of the invagination forms the 
_ommatidia. The more recent work of Giinther (1912) on 
Dytiscus marginalis supports Weismann’s original account. 
Lowne (1893-1895) partly accepts Weismann’s views, but 
disagrees with him in certain important points, in which, 
however, he is undoubtedly incorrect. In support of his 
Dioptron Theory of Insect Vision he wholly denies the pere- 
tration of the fenestrate membrane by nerves; but there can 
be no doubt as to its occurrence in Nasonia. His view of the 
origin of the rhabdomes is very remarkable; he regards these 
structures as arising from the mesoderm and developing in a 
manner analogous to that of the tracheae of the eye (thesé 
_ are highly developed in the blow-fly) ; the perioptic membrane 
he regards as growing out from the brain, although the occur- 
rence of so much neuroglia tissue in that organ has not been 
demonstrated. An examination of fig. 71 (p. 546) of his work 
shows the perioptic membrane communicating with the ecto- 
_ derm on either side of the optic disc; the cells stand off from 
_ the fenestrate membrane, and nerve cells are seen migrating 
into the fibres of the optic stalk, which may possibly have 
been formed from the cells of the perioptic membrane. In 
fig. 6, pl. xxxviii. (p. 548), he actually shows a branching 
a "cell of the perioptic membrane attaching itself to the fene- 
strate membrane at the base of the ommatidia, exactly as I 
ie have described it above in Wasonia. There seems, then, to 


ak teed ae 


| 


384 


be little doubt that the origin of this membrane in Calliphora 
is identical with what happens in this wasp. 

The Ocelli.—The development of these structures can be 
followed from the earliest larvae right throughout larval and 
pupal life to the mature condition of the adult wasp. This 
is rendered possible by the fact. that around those three small 
areas of the head ectoderm from which the ocelli will later 
develop the somatopleure of the head is deflected downwards 
(fig. 66) and becomes continuous with the splanchnopleure 
covering the brain, these curious structures are doubtless the 
remnants of what must once have been a very extensive con- 
nection between the ectoderm and the nerve cord as it sank 


inwards in the embryo. From this fortunate occurrence, it is © 


possible to trace the development of the complex ocellus from 
a stage in which it is represented by a single pair of minute 
_ cells (fig. 66), a condition in which we see it in the larva of 
the first instar. | 

The ectodermal cells covering the head at this stage are 
small in number though rather large and irregular; two cells, 
however, included in each area covered by the conical deflected 
somatopleure are considerably smaller than these. During 
larval life these cells undergo division, so that in the larva 
which is about to defaecate, one sees a conical mass of about 
a dozen cells, rather slender and elongated, in the place which 
in the early larva was occupied by only two (fig. 77). 

These cells continue to multiply mitotically, so that in 
the freshly formed pupa the ocelli are represented each by a 
rounded thickening of the ectoderm in which the cells are 
beginning to arrange themselves in céncentric layers, at the 
same time increasing somewhat in length (fig. 67). During 
the next four hours there is an active proliferation and elonga- 
tion of these cells, giving the whole structure ah appearance 


very like that of a mammalian taste bud. The cells are 


elongated and spindle-shaped, and present each a short pro- 
cess externally. ‘These cells become the visual cells of the 
ocellus, and their short processes, which together form a small 
group at the extremity of the sense organ, project freely 
from it. 

Meanwhile the head ectoderm surrounding the ocelli pro- 
liferates and begins to encroach upon the area which has till 
now been occupied by the ocellar cells. In the twelve-hour 
pupa (fig. 68) this can be seen to result in a gradual con- 
striction of the upper end of the ocellus, which at the same 
time begins to be forced down below the surface. 

In the thirty-six hour pupa this process is complete; the 
ectoderm has grown right across the ocellus, and in its middle 
is seen to undergo a distinct lens-like swelling (fig. 69). 


] 
| 
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>> —— <> “e———- ea 


385 


This growth inwards of the ectoderm surrounding the 
-ocelli not only results in a sinking downwards of the ocellus, 
but it also brings about an almost total closure of the cup, 
_and the cells which now comprise the ocellus are of two kinds, 
the upper ones forming the ‘‘rim’’ and the ‘‘lid’”’ of the cup 
-are small and cubical; they will develop later into the sides 
and part of the “‘iris’’ of the ocellus. The others are the 
developing visual cells; those in the lower part of the ocellus 
‘sink downwards a little, the combined result of these processes 
being to form a cavity in the upper part of the ocellus, into 
which the visual cells project. These cells have meanwhile 
‘become distinctly conical by the broadening out of their bases, 
and their inner ends are beginning to turn towards the 
‘pupil’ of the ocellus, z.e., to the space resulting from the 
Becomplete closure of the distal portion. The nuclei are 
i situated towards the base of these cone-shaped cells. At this, 
_ stage also the distal terminations of the visual cells are begin- 
ning to constrict considerably more, 7.¢., the visual rods have 
commenced to develop. 

; Meanwhile the cells of the ectoderm covering the ocellus 
become irregular. Their nuclei move into their basal portions, 
and the distal ends begin to secrete an outwardly convex 


cuticle. This is the beginning of the ocellar lens (figs. 69, 70). 
i The cells now continue to grow in size, especially the 
“visual cells, which become rather long and robust, with large 
“Prominent nuclei in their basal portions. In the pupa of the 
/ third day the visual rods are completely developed; each has 
apparently been produced by a constriction of the distal por- 
‘tion of the visual cell. These cells also begin to undergo 
Pigmentation at this stage, the pigment granules being con- 
ned to the distal portion of the cell, immediately adjacent 
' to the visual rod. 

o The most obvious features of the development of the visual 
cells at this stage is their marked increase in length, which 
' now results in a considerable lengthening of the whole ocellus ; 
this also appears to bring about a slight downward movement 
‘of the ocellus as a whole, resulting in the cubical cells of the 
distal end assuming a more peripheral position. Meanwhile 
‘the superficial ectodermal cells continue to secrete the lens, 
) which has in the three-day pupa become distinctly biconvex 
(fig. 70). The nuclei of these cells retain their position at 
the base of the respective cells, while the distal end appears 
| to undergo a fibrous degeneration, a change evidently con- 
nected with the development of the lens; so far as I could 
observe, the basal portions of these cells do not. disappear, 
‘but aid the cubical cells in the distal part of the ocellus to 
. the “‘iris’’ (fig. 70). 


“* 
| 


In the four and a half-day pupa these changes are com- 
plete. The lens is strongly developed, biconvex, with the 
greater convexity turned inwards; pigmentation has in- 
creased, and the iris is so disposed as to leave a rather large 
“pupil” space, towards which the visual rods all point (fig. 
71). An isolated adult visual cell is seen in fig. 72. It 
measures about 24 in length, of which 10°4u is occupied 
by the visual rod. ] 


This description holds for the median as well as the lateral 
ocelli; a transverse section of the former, however, shows that 
it is strongly indented in its anterior wall. It is essentially 
a double ocellus; and its double nature can be recognized 
throughout development. As is to be expected, its nerve 
communicates with two ocellar ganglia. 

During this process the mesodermic somatopleure lining | 
the base of the ocellus has grown considerably, and occa- | 
sionally is very prominent. It is retained throughout pupal 
life and is seen in the newly hatched wasp as a distinct mem- 
brane, with fairly prominent nuclei, close to the ocellar wall. 


The innervation of the ocellus is quite different from that 
of the compound eye. In the late larval stages, after defae- 
cation has taken place, nerve fibres grow out from the brain, 
towards the ocelli, guided apparently by the deflected somato- 
pleure. In the newly formed pupa the nerve has already 
come into contact with the developing visual cells of the 
ocelilus, and the only visible change undergone by the nerve 
as development advances is an increase in size; in the early 
pupa it is long and slender, but as the ocellus is forced below 
the surface of the head, and as the brain increases in size, the 
nerve becomes shorter and thicker. It is not necessary to 
describe the formation of the ocellar nerves more fully at 
this stage beyond mentioning that the ocellar nerve is a true | 
nerve, 2.€., quite devoid of cortical brain cells, and therefore 
quite different in nature from the optic ‘‘nerve,’’ which is } 
essentially an outgrowth of the brain cortex. 


386 


C.—Tue Resprratory System. 
The Larval Organs. 


In the newly hatched larva (fig. 1) there is a pair of | 
great longitudinal tracheal trunks passing from the second 
segment backwards on either side of the body to the twelfth 
segment; these are connected with one another, in front and | 
behind, by two transverse tracheal vessels, of which the §} 
anterior passes over the oesophagus, the posterior under the 
rectum. The anterior transverse vessel shows a small, ¥ 
forwardly projecting median part, evidently indicating the 


\j 


387 


point of fusion of the tracheal trunks as they grew inwards, 
_ towards each other, in the embryo. 
y. The longitudinal vessels open to the exterior by four 
_ pairs of spiracles; one on the third segment, a second on the 
' fifth, the next on the sixth, and the last on the seventh seg- 
_ ment, each connected by a rather short stigmatic trunk to 
the great longitudinal vessels of the larva. The tracheal 
_ vessels in the region of the fourth, and the eighth to the 
_ eleventh segments, are provided in each segment with a pair 
of small rudimentary trunks, which are related to certain 
spiracles which do not develop till the next larval moult; 
there are, therefore, nine pairs of spiracles functioning either 
throughout larval life, or only in the later larval instars. 
Besides these nine spiracles there is a pair of tenth 
“rudimentary stigmatic trunks, situated in the twelfth seg- 
ment. They do not open on to the surface till at the time 
of the last larval moult, and become the posterior spiracles 
of the abdomen of the imago. Of the ten potential stigmatic 
_ trunks only nine, therefore, function at some time or other 
_ during larval life. It is interesting to notice that the larva 
of the honey-bee develops the full set of ten spiracles. 
In each segment the tracheal trunks give off a number 
of thinner branching - vessels, which on account of their 
different structure I shall speak of as tracheoles. They are 
clearly seen (figs. 1, 2) in living larvae as fine silvery lines 
ramifying among the organs of the larva; generally there are 
two or three pairs in each segment which pass vertically, 
_as well as, especially in the more posterior segments, several 
_ pairs which run dorsally, but are usually more difficult to see. 
‘The anterior transverse vessel supplies the head by means 
of two groups of tracheoles, which run forwards, but are not, 
at this early stage, very strongly developed. From the pos- 
_ terior transverse vessel several small branches are given off 
_ to the anal segments. 
Structurally there is a very pronounced difference between 

the great longitudinal and transverse tracheal trunks and the 
_ stigmatic trunks, on the one hand, and the smaller branching 
aeracheoles on the other. 
ip In very young larvae the true tracheae are tubes with 
an epithelium of rather thick cubical clear cells, which have 
already secreted the ‘‘spiral’’ intima; before the end of the 
- first instar these cells have become slightly granular, and the 
general growth of the larva is accompanied by a gradual 
‘flattening out of these cells. The stigmatic trunks are similar 
in structure (fig. 93); the spiracles are small cup-like struc- 
| sae lined by an intima devoid of ‘‘spirals.’’ Their intima 
is shed, and a new one reformed at each moult. 


— 


388 


The tracheoles are structurally quite different (figs. 76, 
81); each group or branching system of tracheoles is essenti- 
ally a unicellular structure, of remarkable dimensions. It 
consists of a large clear tube, which soon branches into two; 
from these branches numerous smaller trunks come off, and 
these ramify amongst the organs of the larva. The tubes are 
entirely devoid of a chitinous intima, spiral or plain, and 
never, so far as I could observe, terminate within the cells of 
any tissue. The nuclei are ‘oval, and very large, measuring 
at the end of the first instar Dy in length, 8p in breadth; 
nucleoli are absent; the chromatin is scattered throughout 
the nuclear space, but two karyosomes are generally present 
(fig. 10). The nucleus is usually situated at, or a little 
beyond, the first point of branching. 

The tracheoles of the imago, which will be referred to 
later, differ somewhat from these larval tracheoles; neverthe- 
less, there is a close similarity between the two, and the 
development of the latter may be inferred from what is 
observed to occur in the development of the former. 


There they are formed invariably as outgrowths from the 
tracheal trunks, and there can be no doubt that it is by this 
method that the larval tracheoles are formed during embryonic 
life. They are to be looked upon as modified tracheal 
epithelium cells, which grew in size and developed into cells 
to which the name Grant Tracheoloblasts may be applied. 
These cells then began, still during embryonic life, to grow 
out from the tracheae and developed the tracheoles from 
themselves, as they grew out. It is difficult to determine 
exactly how this happened; probably the great tracheoloblast 
which already enclosed, on account of its size, a considerable 
portion of the lumen of the trachea, began to grow outwards 
at one end. As it grew outwards its two free edges fused 
together, forming a tube. A tracheoloblast in this condition 
leaving the renovated longitudinal vessels and growing out- 
wards to form a tracheole of the imago is shown in fig. 82. 
The cell with its great nucleus then grew further and further 
out, secreting the main branch of the system after it, as it 
advanced. Soon, however, the nucleus ceased to advance, 
perhaps on account of the pressure of the fat body, which 
occupies so much of the haemocoele, and the ramifigation of 
the tubes began by a different method : protoplasmic out- 
growths were produced from the termination of the tracheolo- 
blast, probably by the frequent division, within the main 
substance of the cell, of its lumen. The probable method of 
branch formation within ‘the main portion of the tracheolo- 
blast is shown diagrammatically in text figures A-F. Several 
pairs of these systems of tracheoles occur in a single segment, 


res 


. 
=. 
= 

zs 


Nit 


n 


ty 


Text figs. A-F. Diagram of the Giant Tracheoloblast in trans- 
verse section, showing successive stages in the formation of 
tracheoles. . 


Text fig. G. Developing spiracle of propodeum from the larva 
shortly before pupation. Note the great ‘‘clip,’’ through which 
passes the lumen of the air tube (st.). Closing of the clip is 
brought about by relaxation of the muscles (mcl.). 

Text fig. H. Diagram of respiratory system of adult wasp: sp., 


spiracle; d.l.a.s., dorso-lateral air sac; v.l.a.s., ventro-lateral air 
sac. 


i} 


390 


and the co-operation of these must produce a very efficient 
respiratory system for so sluggish a larva. 

So far as I am aware, no respiratory vessels similar to 
those here described have been observed in other insects. The 
complexity of shape of these great branching cells (fig. 76), 
indeed, finds no parallel, except amongst the nerve cells of 
higher animals. In some ways, indeed, they closely resemble 
these ; and their method of extension is very similar to that 
observed by Ross Harrison in his well-known work on the 
growth of embryonic nerve fibres in plasma media; while as 
an example of a Trophospongial cell, in the sense in which 
Holmgren employs it, they are quite unrivalled. 

In the majority of insects the smaller air tubes are true 
multicellular tracheae, the terminal portion of which alone is 
devoid of spirals and is evidently intracellular; it would seem, 
then, that the great unicellular tracheoles above described are 
homologous with the terminal portion of the tracheoles of 
other insects. Indeed, Pérez (1910, p. 191) gives an account 
of the development in Calliphora of the terminations of the 
tracheoles among the muscles of flight, which is not unlike 
the process by which the large tracheoles of NVasoma are 
developed. From this it would seem to follow that the 
dorsolateral air-sacs of the adult Vasonia, as well as some of 
the great head and abdominal vessels which develop during 
pupal life (see below) are homologous with the general system 
of smaller tracheal vessels occurring in other insects. 

The main change which the tracheal system undergoes 
in the first instar is a slight increase in the complexity of 
the tracheoles; towards the end of this period those stigmatic 
trunks which have not yet opened on to the surface (the 
fourth, and the eighth to the eleventh) grow outwards, and 
at the next moult begin to function. 

The tracheal system has now attained to its mature 
condition, and during the rest of larval life is characterized 
mainly by a considerable increase in size and the extent of 
its ramification, as the larva itself grows. Especially marked 
is this tracheal proliferation in the head region, where the 
brain is developing. The increase in complexity of the 
respiratory system is shown by comparing figs. 1 and 2, 
and is due entirely, so far as I could observe, to an increase 
in the size and complexity of the great branching cells, not 
to a formation of new ones. 

The extensive branching of the tracheoles makes it im- 
possible to measure the size of these, but that the increase in 
bulk is very large is unquestionable. The nuclei of the tracheoles 
show only a slight increase in size. Thus, while the nucleus 
of the tracheoloblast measured 21 by 8 at the end of the 


391 


first instar, at the end of the last it measured 24p by Qu. 
Within thé main tracheae the growth of cell and nuclear 


. size is more easy to estimate. Throughout larval life the cells 


become gradually more granular in appearance, and at the 
end of larval life appear as large flat discs upon the surface 
of the tracheal intima, which has itself stretched consider- 
ably. The nuclei have become greatly hypertrophied; their 
chromatin has become scattered through the nucleus, the 


_karyosomes having disappeared; in their place is found a 


great nucleolus (figs. 84, 85, 86, 87, 92). 

At the end of the first instar the cells of the tracheal 
epithelium measured, on the average, 14 in length, con- 
siderably less in breadth. In the adult larva they measured 
about 34y in greatest length, 10 to lly in breadth. So far as 
I could observe, this great hypertrophy of the nucleus is not 
accompanied by a corresponding increase in the quantity of 
chromatic material; the nucleus extends not in the volume of 
its contents, but by a loosening of its texture. The respira- 
tory system of the mature larva, like the other purely larval 


organs, is to be looked upon merely as a greatly hypertrophied . 


condition of the tracheal system of the newly formed larva ; 
no differentiation of essentially new structures ever occurs. 

Having remained in this condition for about a day (rest- 
ing period of the larva), the tracheal system begins, at the 
time of defaecation and in the post-defaecation period, to 
disintegrate, and by the time the pupa has been formed (one 
day later), only a few disintegrating vestiges of the old 
tracheal system are recognizable. 


The Destruction of the Larval Tracheal System. 


The processes of disintegration of the old larval respira- 
tory system and the regeneration of the systemeof the adult 
are contemporary ; indeed, the imaginal cells often push the 
worn-out larval cells aside, before the leucocytes have had 
time to remove them. Nevertheless, it will be better to con- 
sider the two processes separately. 


The epithelium of the main tracheal vessels begins to | 


disintegrate, at the time of defaecation, and in eight hours’ 


time has wholly disappeared. Besides the presence of the | 


great nucleolus, and a general hypertrophied condition of the ~ 


cells, these show no abnormal characteristics. Occasionally, 
however, distinct vacuolation of the cytoplasm can be observed. 

At the time of defaecation, however, these cells begin to 
suffer attack from leucocytes; this is especially well seen in 
the main longitudinal vessels at about the time of defaecation 
(figs. 83, 86). The actual process of histolysis is difficult to 
observe on account of the smallness of the objects dealt with; 
but that the leucocytes play a large part in the removal of 


Sf 


392 


the tracheal epithelium is clear. The tracheal intima does 
not suffer any corresponding change. 

The destruction of those lateral stigmatic trunks which ° 
do not persist in the adult wasp begins in the freshly formed 
pupa. Here the cells lining the lateral stigmatic trunks 
undergo cytoplasmic degeneration. This stage is easily recog- 
nized on account of the great hypertrophy of the nucleoli, a 
condition so characteristic of the worn-out larval cells of 
Nasonia. In close connection with the disintegrating stig- 
matic trunks leucocytes may occasionally be seen, actively 
removing the débris. Whether the remains of the cells 
(nucleus and cell wall) disintegrate of their own accord, or 
whether leucocytes remove them, I am not definitely able to 
say; the appearance of preparations rather suggested the 
latter. 

Composing the epithelium of the stigmatic trunks are two 
kinds of cells. There are large, purely larval cells, and much 
smaller imaginal cells; it is only the former that grow during 
larval life, and degenerate at the end of it. The imaginal 
cells are clearly seen in even the youngest larvae at the bases 
of the trunks (fig. 93). It is from these ‘‘imaginal nests’’ 
that the whole tracheal system of the imago becomes formed. 
(See below.) 

The whole process of disintegration of lateral spiracles 
occurs in the early stages of pupal life, much later, therefore, 
than that of the tracheae; the stigmatic trunks which dis- 
appear in this manner are the second, the third, and the fifth 
to ninth, only the stigmata of the pronotum and propodeum, 
and the newly formed pair of the twelfth segment (see below) 
being retained. 

The whole system of tracheoles also disappears; in the 
living insect, however, in which the tracheoles are clearly seen 
through the transparent cuticle, no discontinuity in the 
general structure of the respiratory system is apparent. This 
is due to the fact that the new tracheal system is forming as 
__the old degenerates (cf. figs. 88, 91). In the sixteen-hour 
pupa the tracheoles still appear quite normal, though greatly 
hypertrophied. But shortly after this leucocytes begin to 
accumulate round the finer tracheoles of the head cavity and 
»« the process of histolysis commences. Sometimes the leucocytes 
. may be observed forsaking their free) life in the_blood-stream ; 
attaching themselves to a branching system of tracheoles they 
begin to crawl over these, and eventually phagocytosis com- 
mences (figs. 88, 90, 91). By the time the larva pupates (six 
to eight hours later) the finer tracheoles of the head have dis- 
appeared, and the larger ones are rapidly undergoing the 
same fate. 


ee 


ali nate 


ee ee 
a ae 


a Shs ey ym eee th emery ew rere s 


393 


Many of the tracheoles, however, appear to undergo 
mainly chemical disintegration; lying, as they often do, 
closely embraced by the great ‘“‘fat cells,” they seem to be 
protected from the action of the leucocytes. Their protoplasm 
becomes finely vacuolated, the lumen disappears, and the tubes 
gradually fragment. ‘This is especially beautifully seen in 
some of the great head tracheoles, which disappear at about 
the time of pupation (fig. 91). The great abdominal tracheoles 
disappear a few hours earlier, also by chemical disintegration ; 
frequently, however, if the pupating larvae are examined, 
rows of leucocytes in the place where the larval tracheoles 
once were, indicate that phagocytosis of the vestiges of the 
tracheoles has, in the end, occurred. Active phagocytosis may 
be observed at times, however (figs. 87, 88), in places, such 
as, for instance, the cavity of the thorax, where they are easily 
accessible to the wandering phagocytes. Several phagocytes 
may apply themselves to the degenerated tracheole, and, dis- | 
solving parts out of it, gradually absorb it. i 

Thus, partly by chemical disintegration and partly by 
phagocytosis, the whole larval respiratory system, with the 


_ exception of the spiral intima of the lateral longitudinal, and 


anterior and posterior transverse vessels disappears within a 
few hours after pupation. 

The intima of the lateral stigmatic trunks is shed during 
moulting. 


The Regeneration of the Tracheal System. 


The regeneration of the imaginal tracheal system has kept 
pace with the destruction of the larval vessels, and takes _ 
place from the ‘‘imaginal nests’ at the bases of the stigmatic | 


trunks, At the end of the resting period of the larva, the | 


cells composing these “‘nests,’’? having lain dormant during 
the feeding period, rise into sudden activity, and proliferating 
greatly (fig. 89), extend as imaginal tracheal histoblasts in- 
wards and along the intima of the great tracheal vessels, 
pushing the epithelial cells which the leucocytes have not 
removed aside as they advance, and taking up a position 
between the larval intima and the epithelial cells from which 
it was secreted (fig. 84). Some twelve hours later the epi- 
thelium of the larval intima has been completely regenerated 
(fig. 85). Those stigmatic trunks which are to persist in the 
imago undergo a similar renovation (fig. 89). In the others 
this does not take place, and they disappear (fig. 92). 

The histoblasts are at first somewhat spindle-shaped as 
they advance, but they soon spread out and form a thin-walled 
tube In close eee the spiral intima. The further 


394 


history of this newly developed tracheal epithelium will be 
considered later. . 

Meanwhile the last abdominal spiracles have developed ; 
not till this time, therefore, is the number of spiracles com- 
plete. They are formed each as a massive down-growth of 
_ very small cells which, passing inwards.and forwards, develop 
a lumen and soon fuse with the main tracheal trunks; a 
spiral intima develops almost immediately. 

In this spiracle the process of the development of the 
spiral intima could be clearly seen. The intima is secreted 
from the walls of the spacious lumen, and, when the surface 
of the cells which are secreting the intima is examined it is 
seen that they present strong ridges which fit exactly into 
the ‘‘spirals’’ of the intima which is being secreted, and just 
as in the markings of the general body surface, so here the 
spirals are merely the secretions formed upon a previously 
protoplasmic ‘‘mould.’’ As the intima thickens the ridges on 
‘the cells gradually straighten out, and the outer portions of 
the intima, which are now secreted, are devoid of spirals. 

It should, perhaps, be pointed out that the intima does 
not possess a true spiral structure, but is simply thrown in- 
ternally into the form of a series of ridges, closely arranged, 
and giving the optical appearance of a spiral. 

In connection with the spiracle of the twelfth abdominal 
segment, and also that of the propodeal (sixth) segment, a 
remarkable structure develops for the closing of its opening 
_ (text fig. G). A large number of cells of the massive 
ingrowth, which gives rise to the spiracle, arrange themselves 
in the form of a minute bent “‘clip,’’ whose arms'enclose the 
spiracle below. From them is secreted a chitinous bent rod, 
the two arms of which, very closely approximated for the 
greater part of their length, meet, and diverging again are 
strongly curved outwards distally; they thus form a complete 
ring round the trachea a short distance from its opening. The 
distal diverging portions are joined by a number of muscle 
fibres. By contracting, they can loosen the arms of the 
chitinous fork, and so bring about opening of the stigma, the 
distal divergence serving as a lever to increase the efficiency 
of the mechanism. This remarkable structure is distinctly 
visible in the adult wasp, if this has been rendered trans- 
parent by caustic soda. During pupal life the chitinisation 
of the spiracle increases, forming the well-marked structure of 
the adult. 

It is necessary to return now to the further development 
of the main tracheal vessels. 

In the larva some twelve hours after defaecation the 
larval tracheal epithelium has been wholly replaced by the 


395 


imaginal histoblasts, which now extend as a new coating right 
along, and in close contact with, the larval tracheal intima 
(fig. 85). 

But already before the epithelium has been completely 
renovated, the histoblasts at the anterior extremity of the 
longitudinal tracheal trunks begin to grow forwards over the 
numerous tracheoles which all open into the main trunks here; 
as many as eight tracheoles may converge towards this. region 
and become enclosed together in the tracheal epithelium as it 
extends forwards. This process, which begins in the larva 
some eight hours after defaecation, advances greatly during 
the next four hours, and, as a result, a distinct tube is formed, 
which encloses the tracheoles, which now appear in a state 
of degeneration. The appearance of the degenerating 
tracheoles has already been described ; the products of degen- 
eration evidently help to nourish the proliferating tracheal 
histoblasts. 

The tracheal trunks, ceasing to extend straight forwards, 
now begin to grow downwards, and in their further extension 
travel quite independently of the tracheoles ; they are seen four 
hours later as two wide channels running vertically down the 
head, parallel with and internal to the great head tracheoles, 
and often separated from these by the great ascending column 
of myoblasts—the developing musculature of the mouth 
appendages (fig. 91). So rapid has been their development 
that already at this period a ‘‘spiral” tracheal intima is partly 
developed. 

Just before the tracheal trunks turn downwards two out- 
growths are formed from them; of these one grows forwards, 
shghtly outwards and upwards, and supplies the anterior, 
dorsal, and lateral regions of the head. The second branch 
grows out from the descending trunk a short distance below 
this dorso-lateral branch, and gives off a great branching 
tracheole into the brain. In the fresh pupa other tracheoles 
begin to grow out from this ‘‘cerebral trachea’’ ; the structure 
and development of the imaginal tracheoles will be described 
later. 

Tracheoles also entered into the developing antennae, while 
from the main tracheal trunks in the defaecating larva other 
tracheoles extend outwards into the legs and wings. 

In the fresh pupa the-great dorso-lateral air sacs begin to 
develop. The new tracheal epithelium just behind the first 
stigmatic trunk on each side begins to grow upwards as a 
slender column of cells. Cell division continues rapidly, and 
the columns extend further upwards, then backwards and 
slightly outwards, growing as a pair of narrow columns of 
cells, already showing a very distinct lumen, along the 


396 


dorso-lateral regions of the thorax. In the two-day pupa they 
fuse again with the main tracheal trunks immediately in front 
of the propodeal stigmatic trunk. 

The tip of the column presents a remarkable frayed 
appearance (fig. 94); this may be a special adaptation to aid 
the column in forcing its way through the surrounding masses 
of fat cells. 

The cell columns meanwhile have begun to differentiate. 
In the growing columns the cells are thick, rather elongated 
in the direction of growth, and present a clear cytoplasm ; 
the lumen of the cell column is narrow and devoid of intima. 
But at about the time of fusion of the posterior end of the 
columns with the main trunks the epithelium gradually 
flattens, transforming the whole structure from a narrow tube 
into a great air sac (fig. 95). The epithelial cells develop a 
granular cytoplasm; while in the four-day pupa they may 
have nucleoli almost as large as those of the degenerating 
larval cells; a ‘‘spiral’’ intima is quickly secreted. 

In the early pupa a number of other tracheae have devel- 
oped from the main longitudinal vessels; especially prominent 
are two ventral downgrowths, from which the tracheoles of 
the wings have been developed. In the early pupa the 
anterior of these is observed as a thick column of more or less 
cubical cells, which in their descent have torn off and dragged 
along portions of the salivary glands as these were under- 
going phagocytic destruction. A very fine example of this 
is shown in fig. 88; the great wing trachea is observed with 
fragments of salivary glands still attached to it; some of 
the tracheal cells are in a state of great activity and are 
growing outwards to form tracheoles; more anteriorly lie the 
larval tracheoles, undergoing disintegration. 

The tracheal trunk to the hind wing is never so prominent. 

While these tracheae and great dorso-lateral air sacs have 
been developing, the main tracheal trunks have undergone a 
similar differentiation. The epithelial cells gradually flatten 
out, and separating from the “‘spiral’’ intima of the larva 
upon which they have been resting for the last three days, 
soon form the two great ventral air sacs. A ‘“‘spiral”’ intima 
is quickly secreted. The size of the air sac depends, of course, 
upon the degree of flattening undergone by the epithelial 
cells. ’ 

In the defaecating larva a third tracheal system develops 
(fig. 86), in the form of a pair of outgrowths, ventrally from 
near the posterior extremities of the main tracheal vessels. 
They grow out very rapidly; an upper branch supplies the 
intestine and neighbouring organs, while the main part of 
the vessel grows downwards and forwards, and undergoing 


397 


special development in the female, ramifies among the muscles 
of the ovipositor (text fig. H). i 
In the pupa of the fourth day a transverse 


co 


commissural’’ 


. trachea is seen uniting the main longitudinal air sacs; I have 


not observed the manner or time of its formation. 


The old ‘‘spiral’’ intima of the larva can still be seen 
lying within the main air sacs; probably it disappears by 
being drawn out through the thoracic spiracles at the last 
ecdysis; it is certainly not present in the adult wasp. 

The adult respiratory system (text fig. H) consists, then, 
of a pair of main tracheal vessels, dilated in the thoracic 
region into the ventral air sacs, and connected by three trans- 
verse vessels—one anterior, another posterior, the third a 
broad channel joining the two air sacs in their mid-region. 
Passing from the anterior to the posterior end of each ventral 
air sac is a great dorso-lateral air sac. From near the pos- 
terior end of the tracheae a pair of vessels pass downwards and 
forwards and supply the abdomen. The head is aerated by 
a pair of great tracheae which pass forwards from the main 
vessels and divide into three great branches in the head. 
From all these great vessels tracheoles are given off. Three 
pairs of stigmatic trunks, two in the alitrunk, the other in 
the abdomen, connect these tubes with the exterior. 


The tracheoles of the adult insect, though essentially 
intracellular structures, are not such remarkable structures 
as we have seen in the larva. Certain cells of the developing 
tracheal tubes do not flatten out when these form an intima; 
on the contrary, they seem to grow in thickness, and then 
migrating from the epithelium grow outwards in various 
directions and ramify among the organs. They are the 
tracheoloblasts, but they never assume abnormally large 
dimensions. As they grow out from the tracheae (fig. 82) 
they leave tubes of varying width behind them (fig. 95); in 
the case of the smaller tracheoles the whole structure may 
remain unicellular. The larger tracheoles, however, such as 
those of the brain or of the appendages, are multicellular 
structures; their formation can be clearly observed in the 
legs. The nuclei of the small tracheoloblasts occasionally 
divide, and the cytoplasm between the two nuclei thus formed 
becomes drawn apart. This process is repeated several times 
and eventually the tracheole is seen as a narrow tube with a 
number of oval thickenings, the nuclei, along its path (fig. 
103). The tracheoles are essentially protoplasmic structures ; 
I could not detect, with any certainty, a chitinous intima. 
Frequently chromatic (?) granules may be observed within 


the walls of the tracheae between the nuclei. 


~ 


398 


The changes, then, which the respiratory system under- 
goes during metamorphosis—a re-development of the longi- 
tudinal trunks, the formation of a new spiracle, development 
of new tracheoles from tracheoloblasts, the production of the 
air sacs from purely embryonic cells, and, finally, the dis- 
appearance of ancestral stigmata—are identical with the 
changes. which have been going on during early larval life 
(and probably during embryonic life, if these were known), 
or which would have gone on had the whole of the develop- 
ment from egg to adult taken place within the egg membrane. 
The significance of this will be discussed in the second part 
_of this paper. 

The destruction of the epithelium of the main tracheal 
vessels by leucocytes has been described by Pérez (1910) in 
Calliphora; but, so far as I am aware, the general disintegra- 
tion and total renovation of the branching vessels have never 
been observed. 

The conclusions of Breed (1903) and of Anglas (1904) 
that tracheoles are not formed as direct outgrowths from the 
main trunks appears to be quite erroneous, and the criticism 
of Poyarkoff, that Anglas was really dealing with myoblasts, 
seems to me entirely justified, since these cells in NVasona 
“frequently show a remarkable resemblance to small nucleated 
tracheoles. 


THE MUSCULAR SYSTEM. 


The history of the muscular system during post-embryonic 
life will best be considered under the following headings :— 
(a) The anatomy of the larval muscular system; (b) the 
structure of the larval muscles and their post-embryonic 
development; (c) disintegration of the larval muscles; (d) 
regeneration of the muscular system. The last section will 
be considered under various headings, viz.- (1) The longi- 
tudinal abdominal muscles; (2) the vertical abdominal 
muscles; (3) the pharyngeal dilators ; (4) the muscles of the 
mouth appendages ; (5) the muscles of the legs; (6) the ovi- 
positor muscles; (7) the great thoracic muscles (muscles of 
flight); (8) the intestinal muscles; (9) the muscle insertions. 
An examination of all these different muscles, moreover, will 
enable a comparison to be made between them. 


The Anatomy of the Larval Muscular System. 
Although the individual muscle fibres of the larva 
undergo a considerable amount of differentiation during larval 
life, yet the general anatomy of the muscular system does 
not alter. I shall describe it here as it can be observed in 
living larvae in the first instar, before they have become too 
gorged with food to be sufficiently transparent for observation. 


399 


The muscular system consists of three prominent sets of 
muscles: the great longitudinal muscles; the great transverse 
(oblique) muscles; and the masticatory muscles, including the 
dilators of the pharynx. 


The longitudinal muscles (fig. 1) are in the form of twenty 
to twenty-two bands of muscles, passing from one end of the 
body to the other. Posteriorly where the body tapers off, they 
all tend to converge towards one point. Anteriorly they are 
inserted upon the walls of the first larval segment; here also 
they converge, but are less concentrated than at the hinder 
end. The muscles are arranged similarly on either side of 
the median line, and are quite absent beneath the ventral 
nerve cord. 


The transverse (oblique) muscles (fig. 1) are in the form 
of nine pairs of muscles stretching from the third to the 
eleventh segments. They pass in hoops round the body of the 
larva, upwards and backwards; their lower and upper inser- 
tions are generally at the junctions of the larval segment with 
the segments immediately before and behind it respectively, 
a.e., the oblique muscles are generally intra-segmental. This 
is not, however, entirely the case, as the last oblique muscle 
is inserted ventrally on the ninth segment, dorsally at the 
posterior limit of the tenth. Others also stretch over more 
than one segment. 


The muscles of feeding are in the form of a number of 
structures which are inserted upon the pharynx at one end, 
while their other extremity is attached to the walls of the 
head, or to a specially thickened cuticular portion of it—the 
tentorlum. They are the dilators of the pharynx. To the 
tentorium are attached also two very minute muscles 
which move the minute jaws. Only one muscle is attached 
to each mandible, the latter evidently swinging backwards, 
after functioning, as a result of the elasticity of the surround- 
ing cuticle. There are six pairs of pharyngeal dilators. Of 
these the lower two are inserted upon the tentorium. Two 
other pairs, attached to the dorsal side of the pharynx, are 
inserted upon the dorsal head cuticle; while two other pairs 
radiate outwards towards the lateral head walls. The united 
pull of these muscles during feeding would dilate the pharynx 
ccnsiderably and would permit efficient sucking of the con- 
tents of the fly pupa, once the mouth was applied to the 
ruptured cuticle. In young larvae the dilators of the pharynx 
exhibit a thick dilatation along a considerable part of one 
side. This swelling becomes less prominent as the larva 
grows, but is recognizable even in adult larvae. Its nature will 
be explained later. 


400 


The Structure and Post-embryonc Development of 
the Larval Muscles. 


The General Body Musculature. 

The nistology of insect muscle can be very clearly observed 
in material derived from Nasonia larvae, and as a number 
of structures, not hitherto observed, were revealed by the 
Haidenhain haematoxylin method of staining employed here, 
I shall briefly describe the structure of muscle fibres, as it 
occurs in this insect. 

The longitudinal body muscles will be considered first. 
The portion of a single muscle band situated in any one seg- 
ment, and inserted at its anterior and posterior extremities, 
is a single muscle fibre, containing three to five nuclei (and 
developed, as will be seen below, from as many cells). The 
inner, posterior portion of one muscle fibre, z.e., of one intra- 
segmental muscle, is connected by a short process with a 
similar process given off from the outer, anterior part of the 
succeeding muscle (fibre) of the longitudinal band immedi- 
ately internal to it (fig. 99), z.e., there is a ‘‘dovetailing’’ of 
muscles (muscle fibres) not unlike what occurs in vertebrate 
cardiac muscle, which results in a direct communication 
between all the longitudinal muscle bands. This ‘‘dovetailing”’ 
is particularly clearly seen in young larvae and in the hind 
region of adult larvae. The connecting piece is always devoid 
of a nucleus (fig. 100), and does not, therefore, represent a 
distinct cell (see below). 

When a muscle fibre is examined in sections, the longi- 
tudinal fibrillae are very clearly seen; each consists of a 
number of minute spindle-shaped sarcomeres (fig. 127), the 
‘‘spindle’’ shape being due to the concentration of the fluid 
contents at its middle. At other times the liquid contents 
pass to either end of the sarcomere, leaving a clear space in 
the middle (Hensen’s line) which may be “quite wide. The 
two dots often figured on the ends of each sarcomere are 
optical representations of the extremities of the sarcomere 
(fig. 127). Moreover, Krause’s membrane is apparently not 
a membrane so far as the muscle fibre is concerned ; though 
the contrary view is sometimes held, it appears to consist 
rather of closely concentrated minute ‘‘dots,” each repre- 
senting the point of junction of successive sarcomeres. Fig. 127 
shows a muscle portion of a fibre in longitudinal section ; 
the individual sarcomeres, each a spindle-shaped structure, 
are clearly visible, and Krause’s membrane is the effect 
obtained by the junction of successive sarcomeres approxim- 
ately along one line. It is only with respect to the fibrils 
apparently that we can speak of a ‘‘Krause’s membrane’’ as 


401 


a membrane. At any rate, Krause’s ‘“membrane’’ is devel- 
oped by the individual fibrils, and, if adjacent ‘“‘membranes” 
do unite, then the structure is secondarily, not primarily, 
a membrane. I shall refer to this again in connection with 
the structure of the adult muscles. 

In connection with the ‘‘transverse’’ striations, a curious 
fact was noticed which has not, apparently, been hitherto 
recorded. The striations do not run transversely across the 
muscle fibre; on the other hand, the fibrillae are so disposed 
that their thickenings in the fibre as a whole are disposed 
in the form of a double smral (fig. 101). This double spiral 
is not always visible in longitudinal sections, as the muscle 
fibre may have been so cut as to show only a portion of it; 
under these circumstances it will appear either as true trans- 
verse striations or, as a single spiral. _ However, in moderately 
thickly-cut sections the double spiral is almost always clearly 
visible. Moreover, it is possible to focus on top of a ‘‘trans- 
‘verse’ striation, and beginning at one end and focussing 
alternately downwards and upwards, to travel right along the 
spiral, and finally arrive at the other end of the fibre. Also, 
after following a spiral striation through a single turn one 
arrives, not at the succeeding striation, but at the second in 
advance, showing the double nature of the spiral. By no 
conceivable bending or twisting of the muscle fibres could 
true transverse striations be thrown into this form, and the 
question of artefacts can be discarded; moreover, the double 
spiral may be detected in entire muscle fibres if these have 
been sufficiently stretched to allow the spiral on the distal 
side of the muscle to show through the thickness of the fibre. 

Krause’s ‘“‘membrane,’’ of course, is likewise disposed in 
a double spiral. 

The sarcomeres of either end of the muscle frequently 
have only one “‘Krause’s membrane’’; the outer end of the 
sarcomere being in this case inserted into the cuticle of the 
larva. Sometimes the fibrils can actually be traced into the 
cuticle, where they spread out a little to procure an extra 
hold (fig. 127). At other times they are inserted on to the 
terminations of integumental cells (fig. 100). The essentially 
integumental origin of the muscle insertions will be referred to 
later, in connection with their development. 

Others of the fibrils, however, do not become inserted into 
the cuticle, but travelling across the border of the segment 
join fibrils from the next muscle of the same longitudinal 
band, forming a very powerful ‘‘Krause’s membrane” at the 
junction (fig. 126). When a muscle is examined in surface 
view these crossing fibrils are clearly seen, giving the muscles 
a particularly frayed appearance at their extremities. 


402 


The muscles are covered with a very prominent sarco- 
lemma. 


The nuclei are three to five in number; they are round 
or oval flat discs; a very large nucleolus is usually present in 
the nuclei of the fully-grown larvae; or two nucleoli may be 
present; a karysome is quite absent and the chromatin is 
scattered in small granules throughout the nucleus (fig. 102a). 
The nuclei, if round, measure about 17» in diameter; if oval, 
19-20u in length, 13-144 in breadth. The breadth of the 
muscle is about. 344; the length varies apparently according 
to the number of cells which entered ihto its formation; on. 
an average the muscles measure about 250, so that the length 
of the individual ‘‘cell’’ composing the syncytium is 63p. 

The oblique muscles possess four nuclei; in their middle 
they show a long slit, mdicating the double origin of this 
part. Their minute structure does not differ from that of 
the longitudinal muscles. ; 

If the muscles of young larvae be examined, the essenti- 
ally multicellular nature of the muscle fibres is clearly seen 
(fig. 100). Three to five cells may be present arranged end 
on end, and the cell boundaries are still unmistakable. In 
the earliest stage (twelve-hour larva) in which I have 
examined them they show distinct longitudinal fibrillation, 
the fibrillae of successive cells in the developing syncytium 
already fusing. Moreover, fibrillae from one fibre have already 
communicated with those of others of the muscle band. 
Striations are just beginning to appear; in some they are 
distinctly visible, in others quite absent. The individual 
cells measure 14u in length, 114» in breadth. The nuclei 
are relatively gigantic and measure 12-14 long by 8-9» broad. 
Nucleoli are quite absent; one or more small karyosomes 
-may be present, but a considerable part of the chromatin 

may be scattered in granules throughout the nucleoplasm 
(fig. 102b). 

Already at this stage, too, the connecting pieces can be 
clearly seen between the adjacent muscle bands. In the 
oblique muscles the four-celled condition is especially clearly 
seen (fig. 99). . 

The muscles grow rapidly; already in the second instar 
the cell limits are scarcely visible. From now on the muscles 
begin to differentiate into the condition in which we see them 
in the adult. The process consists mainly in a special develop- 
ment of the striations, and a general ‘‘loosening’’: of the 
texture of the whole muscle by the development, apparently, 
of more interstitial substance. The nuclei grow considerably 
in size, but the actual chromatin does not appear to imerease 


403 


in quantity ; karyosomes disappear and the nucleoli develop in 
their place (fig. 102a and b). | 

Although the adult muscles may be inserted upon the 
cuticle directly, yet there can be no doubt that such insertions 
were originally integumental cells. Leydig (1885) first put 
forward this view, and it is held by Duboscq (1898), Hennequy 
(1906), Janet (1907), and Pérez (1910). Others have regarded 
the fibrillae as fusing directly with the cuticle, but this view 
seems scarcely tenable. 

If the muscles of young larvae be teased out, it is fre- 
quently possible to observe the fibrils of the longitudinal 
muscles communicating with the cytoplasm of a-long process 
from the flat integumental cells (fig. 100). These processes 
show a considerable degree of chitinisation, and may appar- 
ently chitinise fully before the end of larval life, thus explain- 
ing the insertion of fibrillae upon a non-protoplasmic surface. 


The Dilators of the Pharynz. 

If a larva in its first instar be examined these muscles 
can be observed in the last stage of development (fig. 47). 
The muscles are formed, probably in late embryonic life, from 
a mass of cells which fuse to form a syncytium. Generally 
about four to five cells combine thus, though in some cases 
as many as fourteen to sixteen (judging by the number of 
nuclei) fuse. J have not examined these muscles earlier than 
half-way through the first instar. At that stage the ‘‘trans- 
verse’ striations are clearly seen, again in the form of spirals. 
The nuclei all become concentrated in one place, and collecting 
a certain amount of cytoplasm round them, form the large 
swellings already mentioned. Hach nucleus has a large 
karyosome. : 

The spiral striations do not extend on to the dilated 
part of the cytoplasm; they are confined to the essentially 
contractile region of the muscle. Along this region fibrilla- 
tion has been taking place, but is not yet, apparently, com- 
plete, for the spirals extend outwards, upon otherwise 
quite undifferentiated protoplasm (see fig. 47, at x). Here, 
then, it seems that the (spiral) striations form first in the 
contractile syncytium, and the longitudinal fibrilliation is 
only secondarily developed. In the general body muscles 
the opposite happens; this appears to be the case also in 
mammalian muscle. — . 

Sometimes a single cell of the syncytium may form a 
number of distinct roots of the muscular portion of the 


insertions. As in the body muscles, the muscle fibres are 


always inserted upon the integumental cells, never directly 
upon the cuticle. 


404 


The Destruction of the Larval Musculature. 


Like the other highly specialized larval organs, the larval 
musculature undergoes total destruction at the end of larval 
life. ’ 

The muscles of the larva do not, however, all disappear 
simultaneously. The pharyngeal dilators disappear first, at 
about the time of larval defaecation. Certain thoracic muscles 
begin to disintegrate several hours later. The abdominal 
muscles persist till a few hours after pupation. 


(1) The Dilators of the Pharynz. 

The disappearance of these muscles is closely associated 

with the development of the pharyngeal muscles of the imago, 
and will be more conveniently described there. 
(2) The disintegration of the wpper three pairs of thoracic 
muscles and of the oblique thoracic muscles is closely con- 
nected with the development of the great vertical and longi- 
tudinal thoracic muscles, and will be considered in connection 
with these. The other thoracic muscles disappear early in 
larval life. I have not, however, carefully examined their 
process of destruction. It is improbable that this should be 
unlike the process as we see it in certain abdominal muscles, 
which I shall here describe carefully. 


(3) The Muscles of the Abdomen. 


Many of the longitudinal muscles of the abdomen of the 
larva are disintegrated by the action of the embryonic cells 
of developing imaginal muscles. The description of these will 
be deferred till the regeneration of the muscular system is 
considered. 

On account of the marked differences shown by the ver- 
tical (oblique) and longitudinal abdominal muscles in their 
mode of disintegration, it will be best to consider them 
separately. 


(a) The Longitudinal Muscles. 


It is mainly in the ventral portion of the abdomen that 
disappearance of the longitudinalb muscles independent of the 
action of developing myoblasts occurs. 

Though the process first becomes marked in freshly formed 
pupae, yet, for a considerable time previous to this, the 
muscles have been undergoing a process of internal degen- 
eration. 

In the larvae at about the time of defaecation the nuclei 
begins to appear abnormal;. they have developed great 
nucleoli, much larger than those usually occurring in nuclei, 
which may be, at this stage, crowded with numerous minute 
highly refractile crystals. Most of the muscles, however, 


405 


appear otherwise quite normal, and are still capable of func- 
tioning, though only very feebly. (The shedding of the last 
larval cuticle is itself brought about by muscular movements 
of the abdomen.) 

Shortly after moulting, however, the muscles lose their 
striations, the substance of the striations spreading itself 
uniformly through the fibrils (the substance of the fibrils 
becoming, in consequence, uniformly heavily staining (fig. 
107). 

The nucleus meanwhile undergoes certain changes; the 
chromatic material of the nucleus may change from a rough 
granulation to a fine chromatic dust, the particles of which 
may be clumped together (fig. 107). This dust may be forced 
into the cytoplasm. It may even scatter itself through the 
substance of the muscle, leaving only the empty nuclear mem- 
brane behind. This occurs in certain of the dorsal abdominal 
muscles whose further disappearance is intimately related with 
developing myoblasts, and is rendered possible by the degen- 
eration of the fibrillae into a loose granular fluid. 

At other times, however, the nuclei remain, to external 
appearances, normal, except for the presence of the great 
nucleolus. 

At about the sixth hour after pupation those muscles 
which have not become penetrated by the myoblasts of devel- 
oping imaginal muscles (see below) are fallen upon by 


leucocytes (fig. 105). 


The leucocytes begin to cluster around the dead muscle — 


fibres and the muscles are rapidly absorbed. “Pérez has ex- 
amined the process fully in Calliphora, and I shall describe it 
only briefly here, referring mainly to the points of difference as 
seen in the two insects. 


When the leucocyte has Be cach aolose to the degen-| 


erated muscle it pushes out a pseudopod which appears to 


dissolve its way ‘through the still unbroken sarcolemma.) | 


Within the muscleit/gradually swells out and drags a certain 
amount of the leucocyte after it; so far as I could observe, uf 
does not entirely enter the fibre (fig. 129). 


The actual Bl Pleo of oe Bat into the tiga ee, 


it has oo sieieedl he cue more Ba cuparons: ones, dissolving 
off small pieces there, which accumulate within the ‘leucocytes, 
in the form of large granules. 

Oceasionaly, however, a much more voracious leucocyte — 
may engulf long strips of muscle substance; so long, indeed, 
may the strips be, that it becomes necessary to bend them 
about to accommodate them within the body of the corpuscle ; 


i. 


406 


sometimes several such strips may be present. Most usually, 
however, the leucocytes remove the muscle tissue in much 
smaller quantities. Pseudopodia, however, are very rarely 
seen; although the absorbed food is frequently contained in 
a vacuole, and although it is possible that during the killing 
of the leucocytes, in making the preparations, pseudopodia 
may have been withdrawn, it is nevertheless quite probable 
that a considerable amount. of feeding takes places by the 
absorption of liquid material, perhaps dissolved by extra- 
cellular enzymes directly through the walls of the leucocytes. 
I shall refer to this again later. 

That the muscles, once their sarcolemma has been rup- 
tured, may undergo a certain amount of ‘‘chemical disintegra- 
tion’’ is not unlikely; it might, however, be very difficult to 
detect microscopically. In the case of the vertical abdominal 
and the pharyngeal muscles, however, it does occur, and is 
fairly easily seen (see below). Nevertheless, the main factor 
in the removal of the degenerated fibre is the phagocytic action 
of the leucocytes. These scavengers, having gorged them- 
selves at the expense of the dead tissue, gradually move to 
some secluded corner in the cavity of the appendages, or 
amongst the developing integumental cells, and there attempt 
to digest their meal in peace. 

The removal of the dead muscles is accomplished within 
several hours; ten hours after pupation they have entirely 
disappeared. . 


(6) The Vertical (oblique) Abdominal Muscles. 


Although myoblasts may, in the anterior part of the 
abdomen, develop in relation with some of the degenerating 
vertical muscles, yet the appearances which these present, as 
they disintegrate, are quite different from those which we see 
in the longitudinal muscles. 

The nuclei present the usual features of a greatly hyper- 
trophied nucleolus, often containing numerous minute 
crystals. The contractile part of the muscles may disintegrate 
at a remarkably early period, viz., in the defaecating larva; 
at other times distinct striations may still be seen a day 
later. Almost invariably, however, the striations have dis- 
appeared from the muscles sixteen hours after defaecation, 
and the resulting appearance of the muscle depends upon 
whether the contractile substance has been cast bodily out 
of the fibre, or whether it has become uniformly scattered 
along the fibrillae. Both these processes occur. I shall first 
describe the former. 

It was in one of the posterior abdominal muscles in the 
larva at. about the time of defaecation that I was able to 


407 


observe the process of disappearance of the striations. The 
muscle is reproduced in fig. 104. In the lower region of the 
muscle the spiral striations are still visible, as heavily staining 
thickenings of the fibrillae, and are still, to all appearances, 
quite normal. In the upper part of the muscle the striations 
have entirely disappeared. In their place the whole muscle 
is filled with a fine dust of disintegrating striated material 
which is being’ thrown in a shower of particles, at first sight 
resembling bacteria, into the blood stream. Some of the 
striations in the individual sarcomeres are still intact, and 
may be arranged apparently quite normally, successively 
along a fibril. Others, already shortening, lie in the inter- 
stitial substance, where they are quickly rounded off, and by 
the time they reach the blood stream, are seen simply as 
minute rounded globules, evidently undergoing solution in 
the blood plasma. The striated substance has not’ been 
pressed out at the end of the fibril; it seems to burst its way 
through in each sarcomere, apparently in the region of junc- 
tion of successive sarcomeres (‘‘Krause’s Membrane’’). 

This loss of material causes a considerable shrinking of 
the contractile substance within its sarcolemma. 

At other times, the muscles do not lose their staining 
reaction; on the contrary, though their striations disappear, 
the fibrillae (?) stain quite strongly, and it is seen that the 
striated substance, instead of forcing its way through the 
muscle sheath, has now spread itself along the fibrillae. The 


latter process appears to be much the commoner of the two. 
_ Exactly what determines which of the two processes should 


occur I am quite unable to say. 

Degeneration of the muscle fibre now continues; the 
sarcoplasm becomes granulated and develops, in_ places, 
rounded globules. Quite frequently these globules absorb 
a granule into their middle, which may give them the appear- 


_ ance of minute nucleated cells. Such a condition has already 


been described in the degeneratmg integumental cells. The 
sarcolemma may have become strongly wrinkled. 

As a result of the degeneration of the interstitial sub- 
stance the fibrillae become pressed close together. Leucocytes 


now penetrate the sarcolemma and a phégocytosis of the inter- 


stitial substance commences. A ‘considerable part of it, how- 
ever, undergoes chemical disintegration, being cast into the 
body cavity as large round globules, which are not to be con- 
fused with globules from the fat-body (fig. 110). — 
_ As a result of this process the fibrillae are set free, and 
falling apart, spread out a little, producing structures of 


_ very characteristic appearance (fig. 110). Sometimes, it would 


seem, several fibrillae cluster together, and the osele is 


noe 


408 


represented not by the loose individual fibrils, but by a 
number of loose bundles, each consisting of a few fibrillae 
which have become fused together. 

Having removed the more palatable interstitial substance 

the leucocytes now turn their attention to the fibrillae. The 
process of destruction seems to be much more difficult here. 
The leucocytes apply themselves round a piece of the fibril, 
and several such leucocytes may often be seen, arranged side 
by side along a single fibrilla in their attempt to destroy it. 
The process is actively going on in the fresh pupa, and six 
hours later the fibrillae have entirely vanished. Muscular 
regeneration often occurs in connection with these degen- 
erating fibres and will be referred to below. 
} The histolytic action of the phagocytes on the muscles 
was first discovered independently by van Rees (1884), and 
by Kowalevsky (1885), using Calliphora as material. 
Korotneff (1892), on the other hand, could not observe it 
in the moth Jznea, but his observations were made on the 
thoracic muscles. He regarded these, apparently erroneously, 
as arising by regeneration of the larval muscles. Berlese’s 
conclusions have already been referred to earlier. 

In Calliphora Pérez finds that the leucocytes generally 
enter the muscles, and break off small pieces of muscle. This 
muscle has not undergone any visible degeneration; even 
_within the leucocytes it seems to retain its structure for a 
considerable period. In Nasonia the degeneration of muscles 
of disorganization Fron muscles hiek have: merely lost or 
even only incompletely lost their striations, to others which 
_ have undergone total granular degeneration, may be observed. 


The Regeneration of the Muscular System. 


An examination of the process of regeneration of the 
muscular system revealed the remarkable fact that a con- 
siderable difference exists in the actual morphology of the 
various muscles of the adult wasp; the muscles of the legs, 
ovipositor, and mouth appendages have similar methods of 
development, and though the mature surface abdominal 
muscles are similar to these, their mode of development is 
quite different. A single pharyngeal dilator muscle, on the 
other hand, corresponds not to a single leg muscle, but to a 
whole group of them; while the great thoracic muscles (wing 
muscles) are quite unique in that they are composed of a 
great many muscle fibres, all running parallel to one another, 
‘ and quite devoid of fibrillae. These remarks will become 
clearer when we have considered the development of the 
various muscles; it is enough to say here that failure to 


* foe? ae 


409 


observe the muscles in their embryonic state has resulted 


in a considerable misinterprotation of the structure of the 
mature organs. 
The adult muscles all arise from mesodermal cells, the 
myoblasts, which are recognizable in the earliest larvae. The 
assertion of de Vaney (1902) that these cells are hypodermal 
in origin, is quite erroneous, and the opinions of Kowalevsky 
(1887), Berlese (1901), Henneguy (1904), Karawaiew 
(1898), Pérez (1910), and finally of Poyarkoff (1910), that 
they are essentially mesodermic cells are easily verified in 
Nasonia. 


(1) The Superficial Longitudinal Abdominal Muscles. 


ee 


rns 


As the most direct development of adult muscles occurs 


in the superficial abdominal muscles, it is best to consider 


these first. 

In the fresh pupa, the longitudinal abdominal muscles 
begin to degenerate. After losing their striations the fibrillae 
cluster togéther in the middle of the muscle, while the inter- 
stitial substance, which in life separates them, becomes forced 
to the periphery of the muscle fibre, appearing here as a 
granular fluid, after showing fatty globules (fig. 107). At 
other times the whole muscle fibre, not merely its interstitial 
substance, may undergo granular degeneration, and the 
chromatic material, breaking out of the nucleus, may scatter 
itself as fine granules amongst the degenerate cytoplasm. The 
sarcolemma remains intact (fig. 108). Some of these muscles 
undergo phagocytic destruction, as above described. It is to 
the remainder that I refer here. 

The myoblasts now become active. During larval life 
these have been lying, as small embryonic cells, 54 to 6m in 
diameter, scattered in the body cavity close to the muscles. 
They now begin to multiply, mitoticallyit seems, Jand, pene- 
trating the sarcolemma, lie in the degenerated muscle 
cytoplasm (figs. 106, 108, 109), where they move about by 
amoeboid action (fic. LOC} 

Within the muscle fibre these cells multiply, and grow at 
the expense of the degenerate larval muscle substance. In 
those muscles where there has occurred a total cytoplasmic 
degeneration their task seems comparatively easy; but in 
those muscles where the fibrillae have failed to disintegrate 
they at first confine their attention to the granular inter- 
stitial substance. Eventually, however, the whole larval 
muscle (fibre) disappears, including even the sarcolemma, and 
the myoblasts are seen in its place. The cytoplasm of the 
myoblasts is always clearly seen, and pseudopodia are often 
visible (fig. 107); but whether the myoblasts absorb the 

M 4 


410 


muscle cell contents phagocytically, or whether they merely 
absorb it through their permeable cell wall, I am unable to 
say. Perhaps, after all, the only function of the pseudopodia 
is to enable the myoblasts to crawl into the position they are 
to_assume in the mature muscle, just as the cells of the 
imaginal integumental areas do (see above). 

It is scarcely necessary to remark that the myoblasts 
in the above account have not been confused with leucocytes, 
the activities of which are not altogether unlike those of the 
myoblasts; leucocytes are considerably larger than myoblasts 
and always have a characteristic nucleus. 

After twelve to fifteen hours the myoblasts of each 
muscle fibre have arranged themselves, one after the other, 
in a row; the pseudopodia have entirely disappeared (fig. 111) 
and the cells are almost cubical in shape. In this condition 
they remain for a long time, the only visible change being 
that they first adopt a very regular arrangement, and in the 
thirty-six hour pupa fuse to form a long columnar syncytium, 
with the nuclei regularly arranged along it from one end to 
the other (cf. fig. 115). But in the middle of the third day 
of pupal life the developing muscle fibre begins to differentiate, 
and first undergoing fibrillation, then striatiom, develops 
eventually into the muscle as we see it in the adult. The 
striations, as usual, are spirally arranged. The nuclei occur 
right in the middle of the fibre (fig. 116).. It follows, of 
course, that a single longitudinal abdominal muscle consists 
of only one fibre. 

The nature of the muscle insertions will be referred to 

later. 
= Pérez (1910) has observed the metamorphosis of the 
abdominal muscles in Calliphora. He finds that the muscle 
fibres lose their striations and fibrillations, and that even the 
sarcolemma is added to the degenerate mass. He described 
the_myoblasts as entering the dead larval muscle fibre 
apparently by amoeboid action. Here they lose their cyto- 
plasm and increase by direct division to form the syncitial 
) mass, which, on differentiating, produces a mature muscle 
fibre. It should be remarked that the myoblasts have only 
an extremely fine pellicle in Vasonia, and that if the degen- 
erate larval cytoplasm is at all compact in consistency, as it 
is in the pharyngeal and thoracic muscles, the myoblast 
cytoplasm is hardly, or not at all, distinguishable, unless, as 
is very often the case, the myoblast lies in a distinct vacuole 
within the mass, part of which it has, apparently, been 
absorbing. The degenerate cytoplasm of the abdominal muscles 
is, however, so loose in texture that the cytoplasm of the myo- 
blast is easily recognizable. The apparent loss of cytoplasm as 


el — le — en <2 Sor oT eS cD —T —- —= Se 


Sacer Ser) eee eT 


411 


described by Pérez, moreover, would be difficult to interpret 
in terms of our usual conception of a living cell. 


(2) The Vertical Abdominal Muscles. 

The development of these muscles takes place in close 
relation with the degeneration of the vertical (oblique) 
abdominal muscles of the larva, which, as described above, 
after losing their cytoplasm, break up into vertical fibrils or 
groups of fibrils. While these fibrils are undergoing phago- 
eytic destruction the myoblasts crawl along some of them and, 
poansng themselves at their expense (fig. 110), ultimately 


BP eitodinal muscles ; unese on |. diitnan Een form fie vertical 
abdominal muscles whose structure does not differ from that 


of other muscles of the abdomen. They are especially well 


developed at the anterior extremity of the abdomen, where 
they become attached in front to certain small phragmas 
within the petiole, and act as the flexor and extensor muscles 
of the abdomen. 

(3) The Dilators of the Pharynz. 

The larva possesses six pairs of pharyngeal dilators 
(muscle fibres) to whose development I have already referred. 
In their neighbourhood, even in the earliest larvae, can be 
seen occasional myoblasts measuring usually some 5u to 6p 
in length. 

Like the other purely larval cells, the pharyngeal dilators 
undergo degeneration, this occurring at the time the larva 
defaecates; but their disintegration differs somewhat from 
that. of the muscle fibres of other_parts-of-the body. The 
nucleus presents the usual ‘hypertrophied appearance, and 
contains the gigantic nucleolus, so characteristic of the degen- 
erating cells. As in the case ‘of the abdominal muscles, the 
pharyngeal dilators, after degeneration, become the prey of 
the_proliferating myoblasts of the imaginal muscles. But 
before they penetrate the muscles these often undergo a 
partial globular degeneration, and these globules, breaking 
through the sarcolemma, are in part cast into the body cavity, 
where they dissolve in the blood (figs. 117, 118, 119). Some- 
times several such globules, floating about in the blood, may 
be gathered up by a leucocyte, if one happens to be present 
(iig...119).. 

Only a portion of the muscle fibre, however, disintegrates 
in this way; some muscle fibres, indeed, hardly change their 
appearance, the only indication of disintegration being the 
refusal of the striations to absorb stains; and between these 
two extremes all conditions of degeneration may be observed 
—from fibres which lose their striations but retain their 

M2 


412 


fibrillation, to others which undergo total disorganization, 
but fail to cast their contents into the body cavity. 

In the larva at about the time of defaecation the myo- 
blasts, which may often be in the form of spindle-shaped 
cells, proliferate rapidly (fig. 117), and an occasional myoblast 
may be observed entering the degenerate muscle. The rupture 
thus made serves for the entrance of the myoblasts, and soon 
several groups of myoblasts, now quite round, may be 
observed, one behind the other, all lying in the path cleared 
by the first myoblast (fig. 121): The cytoplasm of each 

myoblast is, contrary to the observations of Pérez, usually 
clearly visible, lying within a clear space which it has 
excavated out of the muscle substance and the former contents 
of which it has apparently absorbed (fig. 121). Frequently, 
however, the cytoplasm of the myoblasts is so similar to that 
of the disintegrated muscle substance that its limits cannot 
be recognized. More and more myoblasts penetrate the 
muscle fibre till, in the larva eight hours later, the whole 
muscle is riddled with embryonic cells; the sarcolemma seems 
to be absorbed also. During the remainder of larval life the 
myoblasts, after absorbing the remnants of the granulated 
larval muscle, arrange themselves in several columns of cells; 
the cells may be slightly spindle-shaped, at other times brick- 
shaped, and each column is to be considered the equivalent 
of one developing muscle fibre such as I have described in the 
abdomen ;,the pharyngeal dilator muscles, in other words, are 
multifibrous structures, of much greater complexity than the 
ordinary abdominal muscles. By this process six pairs of 


“pharyngeal muscles of the adult are laid down; two other 


pairs are developed from myoblasts which appear to grow 
quite independently of the larval muscles. At any rate, eight 
pairs of muscles are to be observed in the newly formed pupa 
(see fig. 154). 

In the fresh pupa the muscles begin to differentiate. 
Each column of cells becomes a long columnar syncytium, 
just as occurs in the abdominal muscles, so that in the fresh 
pupa the developing muscle consists of a number of syncytial 
columns packed close together (fig. 122). Each column then 
undergoes longitudinal fibrillation, and the whole muscle, 
losing all indication of the individual columns, becomes a 


_ uniform mass of longitudinal fibrillae. ‘The whole process 


goes on very rapidly, and all stages from a non-syncytial mass 
to a true fibrillated mass can be observed in the fresh pupa. 
Even at this time distinct indications of striations can be 
observed, each fibril breaking up into alternate elements, one 
of which stains feebly with haematoxylin, the other with 
eosin. No distinct Krause’s membrane in the individual 


{ 418 


fibrils can yet be observed. Sometimes the muscle fibrillae, 
even before losing their intra-columnar grouping, may show 
_ indications of striations. Sometimes muscle fibres may even 
be observed, one end of which has undergone striation, while 
at the other end striations have not yet developed (fig. 123). 
The visible changes in the development of the contractile 
part of these muscles during the rest of pupal life consists 
in a greater strengthening of the striations and the develop- 
ment of Krause’s membrane. 
| Meanwhile the nuclei have moved from within the muscle 
to the surface, where they lie often in quite prominent masses 
of uncontractile cytoplasm (fig. 123). The interstitial sub- 
stance of the muscle fibres seems to be produced by the only 
partial fibrillation of the syncytical columns. 4, 
The outer walls of the fused mass of myoblasts remain 
as the sarcolemma. 
The development of the muscle insertions is quite simple. 
Each syncytial column, before fibrillating, fuses with a pro- 
cess, several of which may be formed, from the adjacent 
integument. In the late larva these processes are quite long, 
but already in the fresh pupa they have begun to retract 
(fig. 122), evidently exerting a pull on the muscles shortly 
before these differentiate. They soon shorten to the thickness 
of the other integumental cells, and during the third and 
fourth day chitinise, giving the muscle insertions the appear- 
ance of being inserted directly on the chitinous exoskeleton. 
By this process the eight pairs of pharyngeal dilators are 
produced (fig. 124). In structure they are intermediate 
between that of the abdominal muscles on the one hand, and 
that of the muscles of the mouth appendages and of the leg 
muscles on the other. It would seem, indeed, that these 
muscles have been evolved from muscles which once resembled 
the pharyngeal dilators. 
The development and structure of the muscles of the 
mouth appendages and legs, and others similar to them, must 
now be considered. 


(4) The Muscles of the Mouth A ppendages. 


The development of these muscles illustrates a mode of 
formation which differs somewhat from that observed in the 
other muscles above described—a method of formation which 
is to be observed also in the muscles of the legs and of the 
ovipositor. 

Even in the earliest larvae scattered embryonic cells, 

_ with clear cytoplasm and large ‘‘vesicular’’ nuclei, may be 
observed in the ventral portion of the head, in the neigh- 
bourhood of the mouth appendages or their imaginal discs. 


414 


They are distinguishable from the leucocytes on account of 
their smaller size (about 6u) and the clearness of their cyto- 
plasm, which is quite devoid of vacuoles. 

During larval life these cells—the myoblasts of the future 
head muscles—proliferate, but do not appreciably change 
their size or appearance. Whether proliferation is confined 
to the last stages of larval life, or whether it occurs gradually 
throughout larval life, or, lastly, whether it occurs only at 
the time of moulting, I have not observed. At the time of 
defaecation, however, the myoblasts have proliferated greatly, 
and (still dividing mitotically) grow upwards and backwards 
behind the brain as two slender columns of cells (figs. 91, 
154); in the larva eight hours later they have crept right 
up the back of the transforming head, and finally reached 
the dorsal surface. The cells in the lower portions of the 
columns have consolidated themselves, and now form a well- 
defined rod. Those at the growing ends are loosely arranged 
and generally long and ‘‘spindle-shaped.’’ Sometimes they 
are exceedingly long, and apparently represent the cells which 
both Breed and Anglas mistook for tracheoblasts. In growing 
upwards they move along, and support themselves upon, the 
degenerate larval tracheoles (fig. 91). 

In the twelve-hour larva these spindle-shaped cells have 
all adopted the shape characteristic of the other cells of the 
columns; further cell proliferation results in a thickening of 
the columns. 

Although the columns have supported themselves, as 
they grew upwards, upon the great larval tracheoles, they 
soon stand quite independent of these. This appears to occur 
at the time when the most dorsal cells have fixed themselves 
to the ectoderm of the apex of the head. In the larva eight 
hours before pupation the columns have become intimately 
associated with these ectodermal cells. The remainder of the 
development of the head muscles is intimately associated with 
that of these muscle insertions. 

In the pupa in the first day of its existence the cells of 
the two great columns have grouped themselves into a number 
of secondary columns, by the regular arrangement of successive 
cells one above the other. There are thus formed, still within 
the limits of the original columns, numerous secondary 
columns each one cell in thickness; each of these columns 
will become a single muscle (fibre) of the head. 

The dorsal extremities of the two columns, it would 
seem, begin to spread out a little and meet the processes from 
adjacent ectodermal cells—the future muscle insertions. 
These are at first quite long, and even in the larva eight 
hours before pupation may be observed converging from 4 


415 


considerable area of the apex of the head (fig. 154), all upon 
the narrow cell column. During the first day of pupal life 
ectodermal cells still more distant—on a great part of the 
posterior, and also lateral, regions of the head—elongating 
considerably, insert their processes upon the secondary cell 
columns. These processes then apparently contract again, 
and the tension exerted by these appears to overcome that 
which holds the secondary columns together ; they break apart, 
and, the ectodermal insertions contracting more and more, 
drag these columns into the positions they are to occupy in 
the adult insect (fig. 114). 

At their lower extremities the spreading out of the cell 
columns is much more limited; they do not encroach upon 
large areas of the ventral portion of the head, but confine 
themselves to the mouth appendages, which have meanwhile 
developed, and in close contact with which they have always 
been. 

Even in the pupa in its third day the cell columns may 
still be observed in this condition. The ectodermal insertions 
have retracted, and evidently exert a considerable tension on 
the columns. These are seen to consist of about eighty cells, 
arranged one behind the other; only the outer cell walls have 
persisted, so that they now form each a syncytial column, 
already visible as such in the thirty-six hour pupa. Each 
nucleus has a distinct karyosome, lying within the slightly 
granular nuclear space. 

During the fourth day of pupal life the muscles begin to 
show striations—again of the spiral type—and the muscle 
passes into its adult condition (fig. 116). The persisting cell 
walls remain as the sarcolemma. 

The labium is provided with a set of powerful muscles 
which have probably been formed from the great cell columns; 
during early pupal life they become inserted on the posterior 
wall of the head, just above the labium. 

In the proximal joints of the antennae, myoblast cells, 
which in the early larva were dragged into ‘the antennae as 
these grew outwards, form, in the defaecating larva, a cell 
column in the basal joint of each antenna. These cell columns, 
growing backwards, meet the lower portions of the integu- 
mental ingrowths which produce the great cephalic phragmas 
already referred to, and spreading out in a number of separate 
columns on these (fig. 43), produce, by a process similar to 
that above described, the muscles of the antennae. It should 
be noted that only the first joint of the antennae is provided 
with muscles. The cephalic phragmas are strengthened by 
the attachment to their posterior surface, of certain of the 
muscles of the mouth appendages. 


416 


(5) The Leg Muscles. 

The essential features of the development of these muscles 
are similar to those observed in the head muscles. They 
need, therefore, to be referred to only briefly here. 

Excepting the tarsal muscles, for the present, the leg 
muscles are in the form generally of two sets in each segment 
of the leg (fig. 16). Of these one pulls the segment which it 
moves in one direction, the other in the opposite (fig. 18). 
Since, moreover, the joints are of such a nature as to limit 
the extent of movement in one of these directions, while 
the muscles are so disposed as to cause only motion in one 
particular direction, for each muscle, it becomes possible to 
speak of the one as a flexor muscle, and its antagonistic one 
as the extensor, the extensor being that one whose activities 
are limited by the peculiar mode of articulation between the 
segments. 

Of these muscles, a pair, the flexor and extensor tarsi, 
are developed in the tibia, and in the femur the correspond- 
ing muscles of the tibia are developed. In the trochanter only 
one muscle, the extensor femoris, is formed (fig. 17). The 
coxa contains the flexor femoris, as well as, apparently, cer- 
tain other muscles (fig. 16). Proximally these muscles are 
all spread out over a large part of the segment, while their 
distal portions converge and are attached to a tough tendon 
fibrillated in structure, which is inserted upon the upper part 
of the next segment. This suggests, of course, a mode of 
development similar to that observed in the head, — the 
two processes are, indeed, very much alike. 

As the hollow leg- dises grow out from the body in the 
late larva they drag a mass of myoblasts, which lie in close 
contact with the leg-discs throughout larval life, after them. 
In the defaecating larva, while the legs are yet very short, 
these have grouped themselves in each segment in opposite 
columns, in the position they are to assume in the adult, 
z.e., we get the rudiments of flexor and extensor muscles. 
As in the head muscles, the myoblast columns, whose cells 
continue to divide mitotically, grow in thickness. In the 
pupa six hours after defaecation, 7.e., earlier than in the 
head muscles, the upper ends of these muscles become dragged 
apart by the integumental cell insertions. By this means 
cell columns, each corresponding to a single component of 
one of the two muscles of each segment, are produced; the 
proximal ends are spread out, the distal insertions remain 
together and become inserted on the tendons. The muscle 
columns form syncytia in the usual way, which, developing 
striations, transform themselves into the muscles as we see them 
in the adult. The outer cell walls persist, of course, as the 
sarcolemma, 


417 


The tendons correspond morphologically to the body 
phragmas. They are formed as columnar ingrowths from the 
integument, and even in the thirty-six hour pupa still have 
an embryonic appearance. 

The tarsus is provided with a long tendon (fig. 46), 
inserted proximally on the great tendon of the tibia, while 
distally it is inserted on the last segment. It is not unlike 
a tracheole in appearance; in each segment it is dilated, this 
portion bearing the nucleus. In the first and fourth segments 
the tendon gives off smaller branches to the walls. Only the 


fifth segment has a muscle, which moves the claws. 


The tendon is formed as an ingrowth of cells in the 
early pupa (six hours old) which extends right along the 
tarsus and fuses with the tendon of the tibia. 


(6) The Muscles of the Ovipositor. 

In the female there is a remarkable development of 
muscles in the ventral part of the abdomen, which extrude 
and hold the ovipositor in position during egg laying. 

From the great phragma at the upper extremity of the 
ovipositor two great systems of muscles pass to the lateral body- 


- walls (fig. 22). From the lower phragmas other great masses 


of muscles pass to the ventral and lateral regions of the 
abdomen and are all so disposed as to hold the ovipositor 
with a maximum rigidity while this is boring its way. through 
the hard shell of the fly pupa in which the insect is ovipositing. 

The mechanism of retraction of the ovipositor 1s very 
simple. As already described, the ovipositor drags down the 
sternal plate of the preceding segment during oviposition in 
the form of a cone. On the sternum a pair of enlarged 
longitudinal abdominal muscles from the petiole are inserted. 
The pull which these exert on the sternal plate forces the 
ovipositor back to its position of rest. 

The structure of these muscles is identical with that of 
the head and leg muscles. Their development is quite similar. 

In the defaecating larva the myoblasts which have 
throughout larval life lain in this region, proliferate rapidly 
by mitosis. In the fresh pupa they form a solid column of 
cells which passes right along the ovipositor; two pairs of 
smaller columns are seert at the sides of this column. These 
columns then break up in the usual way, being dragged into. 
position by the adjacent integumental cells. Spiral stria- 
tions appear as usual. 


(7) The Muscles of Flight. 
These are the most remarkable, and at the same time 
morphologically the least understood, of the insect muscles. 


418 


Their transformation during metamorphosis has been studied 
more frequently than that of any other organ; nevertheless 
our knowledge of the process, in spite even of the recent 
work of Pérez, is far from correct. Even the name ‘‘wing 
muscles,’’ by which they are generally known, is inaccurate; 
though their function is to move the wings, they usually 
have no direct attachment to these. Most of the observa- 
tions have been made on the blow-fly, Calliphora, and it is 
therefore possible to compare the observations of the various 
authors. 

Kowalevsky (1885) regarded the larval thoracic muscles 
as undergoing phagocytic destruction along with the other 
specialized larval organs; the imaginal muscles. he regarded 
as being rebuilt from a number of mesenchyme cells, lying 
free within the body cavity. 

Van Rees (1889) observed that three of the longitudinal 
thoracic muscles did not disappear; on the contrary, their 
nuclei, he believed, ee Oe while the 
cytes. The newly et ara heeanne spherical, and 
migrating into the muscle substance, transformed this into 
the muscle as it occurs in the adult. 

Korotneff (1892), working with a moth (Tinea) con- 
cluded that the mesenchyme cells found by Kowalevsky were 
superfluous structures in that insect. He could find no trace 
of them in the moth, and believed he could confirm Van Rees’ 
observation on the rejuvenescence of muscle nuclei. The con- 
tractile part of the muscle fibres, as a result of constant 
functioning during larval life, became exhausted, and under- 
went granular degeneration, forming long plasmatic columns 
(‘‘Plasmastrang’’). The rejuvenated nuclei penetrated into 
the mass and formed a separate nuclear column (‘‘Kern- 
strang”’). These gradually reorganized the disintegrated 
myoplasm, and eventually formed the adult muscles. Leuco- 
cytes took no part in the transformation. 

Pérez (1910) re-examined the metamorphosis of the 
thoracic muscles of Callaphora; while confirming the observa- 
tions of Van Rees and of Korotneff that certain larval thoracic 
muscles did not disappear, he attributed to these quite an 
insignificant function in the rebuilding of the adult ae 
‘muscles. |He regarded them merely as the ‘‘scaffolding”’ 
‘which the imaginal muscles arranged themselves. To the 
|“rejuvenated nuclei’? of Van Rees and Korotneff he 
attributed quite a different origin. They were the mesenchyme 
cells of Kowalevsky, and bore no relation whatever to the 
larval nuclei. These myoblasts, as he now called them, 


ae 


em 


Bee eS ee ee eee eee 
- ee : ae 


419 


migrated into the degenerate muscle mass, lost their true cell 
walls, ‘apparently, and growing at the expense of the larval 
muscles, and probably nourished also by the surrounding 
blood, formed a great syncytium, on the outside of which the 
nuclei then arranged themselves. This syncytial mass then 
broke up into five longitudinal masses, and the further break- 
ing up of these masses into longitudinal fibrillae led to the 
formation of structures which, on further differentiation, 
became the adult muscles of flight. 

These conclusions of Pérez are undoubtedly more in 
harmony with our conception of the nature of cells, and in 
the main I have been able to verify them; his observations, 
however, on the differentiation of the great syncytial masses 
are, I believe, quite incorrect; the process as I have seen it 
in Nasoma is certainly entirely different. 

Special attention has been drawn to the flying muscles 
of insects by Schafer (1891), who has formulated a theory of. 
muscle contraction on the basis of certain structural arrange- 
ments which he observed in the ‘‘fibrillae’ of the flying ' 
muscles of insects. The development of the great thoracic 
muscles of Vasoma shows that in homologising the ‘‘fibrillae’’ 
of the flying muscles of the insect with the fibrillae of other 
muscles, Professor Schafer was incorrect ;©) at the same time, 
the false homology will in no way discredit his conception 
of muscle action. The thoracic muscles of insects, indeed, 
are perfectly unique, and deserve to rank, I believe, with the 
other types of contractile structures—plain, striated, and 
cardiac muscles—as a fourth type of muscle fibre, in which, 
though striations are present, fibrillae are entirely absent. 
The flying muscles of the insect are to be regarded not as 
consisting of a great number of fibrillae with remarkably com- 
structure, but rather of great numbers of fibres (sarco- 

es) in. which fibrillae are absent. This will become clearer 

TS we have considered the development of the imaginal 
thoracic muscles. 


The great thoracic muscles of Nasonza are in the form of 
five pairs of longitudinal muscles, lying one above the other, 
and occupying the greater part of the thorax. Anteriorly 
they are inserted upon the ingrown extremity of the meso- 
thorax, while behind they are attached to more or less 


strongly developed phragmas in the region of the metathorax 
and the propodeum. There are, besides these, five pairs of 


(5) A difference between these fibrillae and those of ordinary 


- muscles was indeed considered in Schiafer’s original paper (1891), 


and they were referred to as sarcostyles. 


420 


sternodorsales, attached to the more anterior portion of the 
mesothorax above, while below they are inserted close to the 
origin of the legs. The muscles.which move the wings have, 
therefore, no direct communication with these; their action 
merely causes changes in the shape of the thorax, changes 
which alter the disposition of certain prominences ‘and depres- 
sions on the thorax at the wing insertion, into which fit other 
depressions and projections from the base of the wing. A dis- 
cussion of the mechanism of flight is beyond the scope of this 
paper. 

The development of these muscles begins in the larva 
at about the time of defaecation. An examination of even 
the earliest larvae will reveal scattered cells, the myoblasts of 
Pérez lying near the muscles, but remaining during larval 
life in an embryonic condition. They are not unlike leuco- 
cytes in appearance; but they are smaller, and do not show 
protoplasmic vacuolation, which is so frequently seen in the 
former. 

But at the time of defaecation the longitudinal thoracic 
muscles show distinct indication of degeneration. Sometimes 
the striations merely, become ill-defined. At other times the 
muscle substance undergoes a total disintegration, breaking 
down into a granular fluid which remains within the un- 
ruptured sarcolemma. This condition is especially well seen 
in the larva four hours later. 


But while the majority of larval muscles disintegrate at 
a later stage, under the phagocytic action, apparently, of the 
leucocytes, a certain number of dorsally situated thoracic 
muscles—three, or sometimes four pairs—become enveloped 
by the myoblasts, which shortly before this have become very 
active. 

Sometimes the myoblasts begin to spread over the muscles 
while these are almost normal in appearance (fig. 113); but 
at other times advanced degeneration is very apparent. 

The proliferation of myoblasts, always by mitosis, is most 
marked at the metathoracic and the rear of the mesothoracic 
segment; from here the myoblasts extend forwards in a 
pair of great columns partly upon, partly independently of 
the larval muscles, drawing the neighbouring longitudinal 
muscles together as they advance (fig. 128); in the larva 
at the end of the period of defaecation they have extended 
right along the mesothoracic muscles (fig. 112), while the . 
myoblasts at this stage are beginning to extend along the © 
prothoracic muscles, which have almost retained their normal 
appearance (figs. 113, 129). Three or four of the dorsal pairs 
of longitudinal muscles of the prothorax and mesothorax are 


421 


therefore concerned indirectly in the reformation of the longi- 
tudinal muscles of the thorax of the imago. 


The myoblasts divide mitotically (fig. 112); they are 


rounded ovoid or hexagonal cells with large “‘vesicular’’ nuclei. 


They measure 54p to 64 in width, and have not appreciably — 


altered in size throughout larval life. hf lri dee {renter 


The myoblasts several hours later begin ‘to penetrate the 


tough sarcolemma and work their way into the degenerate 
muscle substance (fig. 129); others follow, and in the larva 
eight hours after defaecation the whole muscle becomes riddled 
with myoblasts, all nourishing themselves, apparently, on 
the-disintegrated larval muscle. They never lose their cyto- 


plasm, such as Pérez describes in Calliphora. 

Any leucocytes which may be close by gorge themselves 
upon the dead muscle (fig. 129), but the myoblasts seem to 
proliferate so rapidly that before the few leucocytes, which 
may be present, have had time to depart they become en- 
tangled in the mass of myoblasts, and rapidly degenerate 
there (fig. 128). They are frequently seen in the early pupa 
—long after the last remnants of the larval muscles have dis- 
appeared—as spherical bodies, a little larger than the 
myoblasts, in which several large heavily staining globules 
are present (y in fig. 135), and in which the leucocyte nucleus 
may still sometimes be observed. The presence of the 
embryonic cells appears to bring about their precocious dis- 
integration. It is these disintegrating leucocytes, I believe, 
that Pérez has taken for the nuclei of the larval muscles, 
undergoing ‘‘chromatolitic disintegration.”’ In Nasonia their 
nature is unmistakable; they do not become apparent till about 
sixteen hours after the larval muscles have disappeared. 

Meanwhile the myoblasts have absorbed more and more 
of the larval muscles, and so extraordinarily rapid is the 
process that in the larva twelve hours after defaecation no 
trace of the larval muscles remains; in their place there occurs 
now a pair of bands of myoblasts lying some distance below 
the integument, on either side of the midline (figs. 130, 132). 


The two columns of myoblasts are at first unexpectedly small, 


measuring only 55y in breadth, 14 to 15» in thickness; they 
extend from the rear of the mesothoracic segment to a con- 
siderable distance into what was the prothoracic segment of 
the larva. 

But already before the end of larval life a series of 
remarkable processes begins, which transforms these two strips 
of embryonic cells into the five pairs of great longitudinal 
thoracic muscles of the imago. In the larva, shortly before 
pupation, certain of the cells of these two myoblastic bands 


422 


arrange themselves in the form of five columns of cells, each 
four cells in thickness. This condition of the bands is seen 
in fig. 131. The cells now appear to lose their inner walls 
so that five syncytial columns are produced, on the periphery 
of which the nuclei are disposed; other cells become incor- 
porated later (see fig. 134). To these columns other myoblasts 
now apply themselves ; these, however, do not become merged 
into the syncytium. On the contrary, retaining their cell 
walls, they may be observed to give off at either end a process 
(fig. 139). These processes grow right along the syncytial 
mass and are the embryonic sarcostyles of which the adult 
muscle contains so many. Shortly after the process can be 
first observed the five syncytial columns begin to show a 
very distinct fibrillated appearance, as more and more myo- 
blasts, applying themselves to the columns, send their fibre- 
like extensions into them. 

The process takes place with considerable rapidity, and 
already in the pupa four hours old the myoblast bands of the 
late larva now consist each of five columns of fibres (sarco- 
styles), surrounding each of which is the layer of cells from 
each of which a single fibre has been formed (fig. 134). 

Between the myoblasts in the early pupa curious heavily 
staining rod-like structures may be observed (x in fig. 135). I 
am unable to say what their significance is. 

The myoblast cells continue to multiply in karyokinesis, 
and the strips grow considerably in breadth and thickness. 

Then, in the pupa about twelve hours of age, the two 
bands at last split up into their five parts, and these are the 
rudiments of the great longitudinal thoracic muscles (wing 
muscles) of the imago. Each muscle consists, at this stage, of 
a great number (between 800 and 900) of fibres, while an actual 
count of the myoblasts surrounding it, really quite a simple 
procedure, showed, in the same muscle, 871 of these to be 
present. This fact, together with the observation of their 
mode of development, can leave no doubt that the muscle 
is built up of great numbers of fibres, each developed from 
one cell, and not of innumerable fibrillae, as is usually sup- 
posed. Between the fibres lies the interstitial substance, 
formed, it would seem, from the five syncytial columns of the 
early myoblastic bands. 

In the thirty-six hour pupa the muscle is still in almost 
the same condition as that just described. The cells have 
now, however, lost most of their walls, and, their outer walls 
alone persisting, these form a structure which is comparable 
with the sarcolemma of other types of muscle. So far as I could 
observe, the connection between the nuclei and sarcostyles, 
which must once have existed, disappears entirely, so that 


423 


from now on each muscle is one great syncytial mass, corre- 
sponding at first sight to an ordinary muscle in constitution, 
but built up, in reality, in an entirely different manner; it is 
a muscle built up of numerous fibres, and not a fibre which 
consists of many fibrillae. 

In the thirty-six hour pupa a very curious thing is 
now seen, the interpretation of which is difficult; each muscle 
now presents a very faint striation, the striations being again 
in the form of a double spiral. The striations never become 
chromatic, as they usually do; nor do they correspond to 
the striations that develop later in the individual fibres. 
It will be seen later, when the muscle insertions are described, 
that these muscles are already pulling on the body wall, 
somewhat as they do in adult life, and perhaps the spiral 
striations represent the direction of the strain within the 
muscle, just as they do in muscles in which they are fully 
differentiated ; but since the ‘‘pull’’ which these muscles exert 
at this time may be of only temporary duration, structural 
differentiation does not follow. It is interesting to note that — 
the distance between successive spirals is almost exactly iden- 
tical with what is seen in other muscles, v7z., about 8n. 

Finally, in the pupa of the fourth day the individual 
fibres (sarcostyles) begin to show transverse striations (fig. 
138); Krause’s membranes, and the striations with Hensen’s 
line between them, are all clearly seen; even the minute 
tubules which Schafer describes appear to be visible; the 
structures, however, are so exceedingly minute that at present 
no further details can be given. The distance between suc- 
cessive Krause’s membranes is about 24u, so that an un- 
differentiated spiral of the thirty-six hour pupa corresponds 
to about three ‘‘striations’ of the component fibres. 

Schafer was unable to observe any clearly defined stria- 
tions in the muscle as a whole; there is no doubt, however, 
that in WVasoma the striations of adjacent fibres are so dis- 
posed as to present a true striation in the muscle as a whole; 
nor is it surprising to find that these striations are disposed 
again in a double spiral. 

Schafer mentioned the occurrence of the nuclei within the 
muscle, 2.€., amongst the constituent fibres; there can be 
no doubt that in NVasonia, and also in Calliphora, according 
to the observations of Pérez, the nuclei surround the muscle. 

The conclusions of Pérez, in regard to the development 
of the imaginal muscles, may be referred to here. This author 
described the myoblasts in Calliphora as spreading themselves 
over five pairs of thoracic muscles. The myoblasts, entering 
these, lose their cytoplasm, and apparently grow at the 


424 


expense of the disintegrated muscle, nourishing themselves, 
perhaps, also upon the blood. At any rate, the syncytium 
formed from the myoblast grows, and then undergoes longi- 
tudinal fibrillation, the ‘‘fibrils’’ corresponding in no way 
with the individual myoblasts. 

The fact that in Masoma the number of fibres is 
approximately equal to that of the myoblasts, and that these 
can, though with difficulty, be observed to form each a fibril, 
renders the conclusions of Pérez in regard to Calliphora 
doubtful; it is also possible that the five pairs of larval muscles 
which persist and into which the myoblasts migrate, are in 
reality the syncytial columns observed in Wasoma, and that 
Pérez failed to observe the earlier state in which larval muscles 
were being overwhelmed. It is, of course, unsafe to argue 
by analogy, but the fact that Van Rees observed only three 
pairs of persisting muscles is significant. 


During pupal life there is a considerable thickening of 
the five pairs of longitudinal wing muscles. In the late larva 
they represent a very narrow strip measuring 55 in breadth, 
14u to 15yu in thickness. In the fresh pupa, when five columns 
of muscles have developed within these, they have become 
more prominent; but they do not yet form the predominating 
organ of the thorax. But in the twenty-one hour pupa, when 
they have broken up into five pairs of muscle columns, they 
begin to replace the fat-body, which has till now filled the 
greater part of the thorax, and growing larger and larger, 
displace this more and more, and with the vertical (dorso- 
sternal) muscles which have meanwhile been developing, 
occupy almost the whole of the cavity of the thorax. 

The development of the dorso-sternal muscles is very 
similar to that of the longitudinal thoracic muscles, and need 
only be briefly referred to here. However, they serve to 
illustrate that the myoblasts may be quite ‘independent of 
the degenerate muscles over which they are extending. In 
the imago five pairs of sterno-dorsales are present; three of 
these are formed by the extension of the myoblasts over the 
three pairs of degenerate vertical (oblique) muscles of the 
three thoracic segments of the larva; but the absence of 
sufficient larval muscles does not prevent the other two pairs 
from developing (the muscles of the propodeum play no part 
in the process, but degenerate in the same curious way as 
do the other vertical abdominal muscles). They simply grow 
as two pairs of vertical cell columns, quite independently of 
any larval muscle. Within each, as also in the other three 
developing sterno-dorsales after the larval muscle has finally 
been absorbed, a single columnar syncytium (not five, as occurs 


el as ee ) 2 es er ial 
- u 


425 


in the longitudinal muscles) is formed by the fusion of a 
column of cells. This syncytium remains as the ‘‘sarco- 
plasm’’ of the future muscle. The other myoblasts then, 
applying themselves to this column, form each a longitudinal 
sarcostyle which, growing along the muscle, eventually 
becomes inserted by its two extremities upon the dorsal: and 
ventral walls of the thorax. Striations, similar to those / 
observed in the longitudinal muscles, occur on the fourth day. ' 


So far as I could observe, the connection between the 
sarcostyles and their nuclei does not persist. The outer walls 
of myoblasts remain as the sarcolemma. 


(8) Intestinal Muscles. 


These are weakly developed ; they will be referred to more 
conveniently in connection with the intestine. 


(9) The Muscle Insertions. 


These are entirely ectodermal cells, which during develop- 
ment force aside the underlying somatopleure and com- 
municate with the developing muscles. 

Sometimes the process is quite simple. The terminal 
myoblasts come into communication with adjacent integu- 
mental cells which now support the muscles. Frequently these 
cells chitinise entirely, or only partly, so that the muscle may 
become inserted directly on the hard chitinous walls of the 
insect. 

Frequently the adjacent integumental cells elongate 
greatly and actively extend towards the developing muscles. 
This may, as we have seen, give rise to the remarkable split- 
ting up of cell columns into individual muscles, such as occurs 
in the head muscles, leg muscles, and muscles of the ovi- 
positor, and there can be no doubt that the integumental 
cells are the active agents which bring this process about. 
In the case of the dilators of the pharynx the cells of the 
muscle insertions are formed from a much more limited area. 
They have the same elongated appearance as have the other 
head-muscle insertions; but there is no lateral pull as these 
processes retract again, and the muscle constituents are not 
pulled apart. It follows, therefore, that each pharyngeal 
dilator muscle is the equivalent of a whole group of head or 
leg muscles, which have all originated by the longitudinal 
splitting of a single column. 

The insertions of the great thoracic muscles are especially 
interesting. The myoblasts at the extremities of the muscle 
columns approach close to the integument (fig. 135). The 
integumental cells begin to divide, in the fresh pupa, trans- 
versely to their length; division is not complete; on the other 


426 


hand, the cells elongate remarkably, producing long threads, 
as much as 75y in length, and consisting each of two or 
three cells joined one behind the other (fig. 136). These 
threads have become inserted into the syncytial columns of 
the developing wing muscles, and as they lengthen, the muscle 
bands contracting a little, become suspended in the upper 
part of the thorax. 

Meanwhile in the newly formed pupa, the ectodermal 
cells of the mid-dorsal region of the propodeal segments 
elongate and grow forwards. Passing underneath the more 
anterior muscle insertions of the metathoracic integument 
they extend forwards and penetrate the muscle column. At 
the anterior end of the muscle columns more distant integu- 
mental cells likewise communicate with the developing mass 
of myoblasts, so that we get a condition not unlike what has 
been observed in the head and leg muscles; the insertion cells 
from a considerable area all converge upon the great muscle 
band. The result when the insertion cells contract is the 
same as what occurs in these other muscles. The two bands 
are pulled apart into their five constituent columns, and these 
on differentiating form the longitudinal thoracic muscles of 
the adult. 

Contractions of the “‘suspension threads’ is preceded by 
a longitudinal splitting of them, and each thread is now a 
unicellular structure (fig. 137). 

The sarcostyles, as they develop within these minute 
columns, communicate several with one thread. The threads in 
the twenty-four hour pupa then begin to contract. The muscles 
become stretched and at the same time the walls of the pro- 
podeum and mesothorax become drawn closer together; in 
this way the arched thorax of the imago is formed. 

The insertion cells meanwhile have secreted at their 
exterior, the cuticle of the integument, but they do not 
undergo complete chitinisation (fig. 137a). On the contrary, 
their more internal parts remain protoplasmic, and split up 
into a number of fibrils, each communicating now with a 
single sarcostyle. In this condition they are seen in the 


imago (fig. 138). 


The Structure of the Adult Muscles. 


From the above description it follows that the adult 
muscles are of several types. The simplest are the longi- 
tudinal abdominal muscles, formed by the fusion of succeed- 
ing myoblasts, in one line. 

The dilators of the pharynx are more complex and 
correspond in reality to a number of these longitudinal 


427 


muscles; they consist of several cell columns, and though in 
| the adult they are not to be distinguished from the abdominal 
muscles, yet in their embryonic state the differences are 
| obvfous. 
The individual muscles which constitute the leg muscles, 
antennal muscles, and the muscles of the mouth appendages 
and of the ovipositor, are similar to the longitudinal 
_ abdominals in structure; but their simplicity is a secondary 
__condition—a whole group of them is morphologically the 
I equivalent of a single pharyngeal dilator. 
ia The contractile substance of all these muscles has assumed 
| the same type of appearance, viz., striations in the form of 
double spirals. In the pharyngeal dilators a certain amount 
of cytoplasm remains non-contractile ; this contains the nuclei, 
sometimes as many as 15 in number, and is frequently in the 
form of a bulging mass on the side of the contractile 
syncytium (fig. 124). In the other muscles the nuclei are 
arranged in a row along the middle of the fibre. 

In their development the fibrillae are first formed; each 
of these then breaks up into successive ‘‘striations’’ as above 
described, adjacent striations disposing themselves in the form 
of a double spiral. The Krause’s membranes are likewise 

. fibrillar structures; but it is not impossible that adjacent 
Krause’s membranes unite, though I have never been able to 
see clear instances of this fusion. It is interesting to note, 
however, that artificial breaks generally occur right across a 
fibre along a series of adjacent Krause’s membranes, indi- 
cating that there is at least some coherence between adjacent 
fibrillae. 

In all cases, the outer cell walls of the syncytium remain 
as the sarcolemma of the muscle. 

But the most interesting of all the muscles are the great 
vertical and horizontal thoracic muscles, and nothing like 
them seems to occur elsewhere among contractile tissues ; they | 


cells co-operating in different ways to form one highly efficient 
organ. The original syncytial mass forms what corresponds_. 
to the sarcoplasm of other muscles, while the contracting fibres 
(sarcostyles), devoid, as they are, of fibrillae, correspond to 
the fibrillae of intracellular structures. It is interesting to 
note that the striations of adjacent sarcostyles are likewise 
disposed in the form of double spirals in the ‘‘muscle’ as a 
whole, and it is particularly suggestive to note, that in this 
case, where the analogy is otherwise so extraordinarily close, 
no connection exists between adjacent ‘‘Krause’s membranes.”’ 
Even the sarcolemma of other muscles is represented, and it 


> 


serve, indeed, as a remarkable instance of a great number of |, 


428 


would be difficult to observe, anywhere, a more beautiful 
instance of a histogenetic convergence—of the development of 
similar structures from embryologically quite distinct elements. 


THE INTESTINE AND RELATED STRUCTURES. 


It will be most convenient to consider these organs under 
the following headings:—(1) The anatomy and structure of 
the adult intestine; (2) the anatomy of the larval intestine; 
(3) the changes which go in the intestine during larval and 
pupal life, and which convert it into that of the imago. This 
will be described under several headings, wz.:—(a) The 
foregut; (6) the midgut; (c) the hindgut. It will then be 
necessary to consider certain closely related structures, w7z. - 
—(4) The salivary glands, and (5) the malpighian tubules. 


(1) The Anatomy and Structure of the Intestine of the Adult. 

The mouth faces downwards and backwards in the posi- 
tion in which the head is usually held. This opens into a 
narrow high buccal cavity continuous above with a dilated 
pharynx, which opens into the oesophagus, a long very narrow 
tube, which passes forwards, then backwards through the 
circumoesophageal nerve ring, and enters the thorax. Here 
it becomes even narrower, and passing through the thorax 
and propodeum as a very fine tube, enters the abdomen, 
where it forms a great dilatation, with fine papery, usually 
collapsed walls—the crop. 

The crop occupies only a small anterior part of the 
abdomen. Behind, it partly envelops and communicates with 
a very short gizzard, which in turn opens into a small drum- 
shaped chamber, considerably shorter than the gizzard. 
Behind this chamber lies the great stomach, occupying about 
one-third the volume of the abdomen, and between the two is 
a structure, formed by the slight projection of the chamber 
into the stomach, which evidently acts as a valve, to prevent 
any forward move of the contents of the stomach. The stomach 
is the true midgut, being endodermal in origin. All the 
structures preceding it constitute the foregut, and are, as 
will be seen, ectodermal in origin. Behind, the stomach is 
continuous with the hindgut, also ectodermal in origin. 

The hindgut 1s composed of two parts, an anterior por- 
tion, the small intestine, and a terminal portion, the rectum, 
which opens in the last segment by a small anus (fig. 22). 
The small intestine does not, however, open into the termina- 
tion of the stomach; it communicates with that organ by a 
small aperture situated on its ventral side about one-quarter 
the length of the stomach from its posterior extremity. The 
small intestine then passes forwards half-way along the 


429 


abdomen, then gradually bending backwards passes as a long 
tube into the rectum. The small intestine is about as wide 
as the gizzard, but in the region of its junction with the 
stomach it is considerably dilated. 

The rectum is a short spacious chamber. Into it pro- 
ject, from its anterior walls, a single pair of remarkable 
organs, the rectal glands (figs. 22, 164). Behind, the chamber 
narrows, and opens by a short duct to the exterior. 

Two structures must be considered in connection with the 
intestine: the salivary glands and the malpighian tubes. 

The salivary gland is in the form of a single rounded 
clump of cells, lying in the postero-ventral region of the head, 
in the midline. It opens by a single duct into the buccal 
cavity. The salivary duct, however, extends far past the 
salivary glands; it travels upwards, along the posterior portion 
of the head, and ends blindly, after making a few irregular 
turns in the anterior portion of the thorax dorsal to the 
intestine. This distal prolongation must evidently serve as a 
salivary receptacle, and itself also contains some gland cells. 

The malyighian tubes are in the form of eight/slender 
thread-like structures bending in various directions and all 
opening into the anterior part of the small intestine, close to 
its opening into the stomach. 


The minute structure of these parts varies considerably. 
The buccal cavity is lined internally by chitin which develops 
numerous thorn-like bristles, all projecting forwards. The 
pharynx is a great dilated portion of this buccal cavity. Its 
walls consist of a single layer of cubical epithelial cells, larger 
behind than in front. The pharynx, as well as all the suc- 
ceeding portion of the foregut, is lined with a thin chitin 
sheath (fig. 124). 

The epithelium of the oesophagus consists in the head- 
and-‘‘neck’’ region of more flattened cells; in the ventral 
portion of the oesophagus before it enters the ‘‘neck’’ there 
is a thickening of this epithelium; and within the thickening 
les a prominent chitinous bar, terminating in front in the 
region of the circumoesophageal connectives, and connected 
behind, by two very short chitin pieces, with the rear of the 
head. The remainder of the oesophagus is a simple fine hair- 
like tube which traverses the thorax close above the nerve 
cord, and enters the crop, within the abdomen. Its walls con- 
sist of extremely minute delicate spindle-shaped cells, with 
their long axes arranged longitudinally (fig. 158). Only with 
the greatest difficulty can nucleus and cytoplasm be observed. 
Internally it is lined by an extremely delicate chitin sheath. 


430 


In this portion of the oesophagus muscles are absent; 
but in the pharynx and the anterior part of the oesophagus 
these are well developed (fig. 124). There are the great 
pharyngeal dilator muscles, whose structure and development 
are described in connection with the general muscular system. 
They are attached by one end to the front walls of the head, 
and behind are inserted upon the epithelium of the front 
walls of the pharynx. Their contraction serves to dilate the 
pharynx. 

Attached to the hind wall are a number of other less 
powerful muscles, which pass upwards and backwards and are 
inserted upon the epithelium of the chitinous thickening on 
the lower side of the oesophagus, above described. 

Besides these muscles there are a number of others, much 
shorter than these, which are distributed longitudinally and 
circularly on the intestine. The longitudinal muscles are 
long spindle-shaped structures forming three or four layers 
on the front of the pharynx; on the oesophagus they are 
much more scanty. The circular muscles are arranged on the 
pharynx in thick bundles, lying outside the longitudinal 
muscles, each bundle being inserted upon the two lateral 
walls of the pharynx, which they partly enclose like a crescent. 
The circular muscles of the oesophagus are thin plates, not 
arranged in thickened bundles. 

These bundles are all of the ‘‘striated’’ type. The 
oesophageal muscles appear to be unicellular. The longi- 
tudinal muscles of the pharynx are composed of five to six 
cells, fused into a syncytium. 

The crop is a curious structure (fig. 159); its walls ne 

of very flat paper-like cells, in which cytoplasm is very muc 
reduced. They closely resemble the cells of the wing epi- 
thelium before this straightens out on emergence, but always 
retain their nucleus and a very small amount of cytoplasm. 
Within their walls lie very fine muscles, which also serve to 
connect them with the crop. 
——~ These muscles are of a type which has not, so far as I 
am aware, been observed hitherto (fig. 125). They contain 
a small nucleus, which lies as a thickening on the fine hair- 
like muscle. But the muscle itself is not of the usual compact 
type, but possesses several branches, which may run in various 
directions ; each branch represents one, or, at any rate, a very 
small number of fibrillae, and presents the usual striations. 
The fibrillae are too limited in number, however, for the 
striations to be able to dispose themselves in spirals, and for 
once it is possible to speak of true transverse striation. 

The gizzard, on the other hand, is a very powerful organ. 
It measures only 70» in length, 40yu in thickness. In shape 


431 


it is roughly prismatic, and is triangular in section. Its walls 
consist of three thickened epithelial plates (fig. 160), each 
bent slightly inwards along its longitudinal median axis. The 
three plates are lined internally by a very tough but elastic 
chitin plate, which envelops them closely. The thickened 
epithelia do not meet along the three angles of the prism, 
and the intestinal epithelium here is much thinner. The 
thick chitin plates are likewise absent here. 

Two sets of muscles are present. There is an inner 
circular which connects the two longitudinal edges of each 
plate. A pull on them will evidently increase the angle at 
which the plate is bent upon its longitudinal axis. Outside 
the circular muscle layer lies a pe of longitudinal fibres. 
All are striated. 

The gizzard is then seen to be a very ingenious con- 
trivance. In section its lumen is triradiate. A contraction 
of the circular muscles causes increased bending of the three 
chitin plates, and they move towards each other and tend to 
close up the lumen. The resulting organ ought therefore to 
prove a very efficient masticatory structure for an insect in 
which feeding is so reduced as in chalcid wasps. 

The drum-shaped chamber immediately behind the gizzard 
is smaller than the latter. Its walls are composed of minute 
cells; these form a thickening several cells deep which pro- 
jects into the stomach and forms the valve referred to above. 


The epithelial cells of the great stomach are large and 
‘““‘brick-shaped’’ in appearance, measuring as long as 23. 
The cytoplasm is granular and vacuolated; the nucleus very 
large and faintly granulated. A great nucleolus may occa- 
sionally be present. A distinct cuticular lining is absent. 

The small intestine (fig. 167) is lined by a single layer 
of irregular elongated columnar cells, each with a very large 
nucleus, and occasionally a great nucleolus; in places its 
epithelium is very irregular, and the chitinous lining formed 


within it presents these same irregularities. The irregularity 


evidently allows of greater distension. There is in places a 
highly developed coat of thick circular muscle fibres; longi- 


. tudinal fibres are also present, being long and spindle- -shaped 


and presenting a thickened nuclear swelling at their middle. 
The fibres are, as usual, striated. 

The epithelium of the dilated rectum (fig. 168) consists 
of very large, rather flattened cells, frequently presenting a 
great nucleolus. The usual chitin lining i is present. The very 
powerful muscle coating is in the form of a single layer of 
broad, flattened, contractile plates presenting the usual 
striations. 


432 


Projecting into the rectum from its anterior wall is a 
single pair of rectal glands (figs. 22, 164). Hach is some- 
what pyramidal in shape, and presents an outer syncytial 
region, covering an inner medullary region in which cell 
boundaries are sometimes just visible. The development of 
the organ, which will be considered later, shows that the outer 
cortex is merely the fused outer ends of the large cells which 
form the medulla and which sometimes lose their individuality. 
Nuclei do not, therefore, occur in the outer zone, but its 
distinctness, especially in immature stages, is very obvious. 
The cytoplasm of the cells.is granular. The nuclei are large 
and granular and have a gigantic nucleolus. At the base of 
the pyramidal mass is a cavity, which is continued as a narrow 
duct upwards through the whole organ. The cavity does not 
appear to open into the body cavity; on the contrary, below 
it the lining of the rectum undergoes a special chitinisation. © 
Through this chitin piece passes a large tracheole, which runs 
through into the rectal gland and there terminates. At the 
base of the pyramid are a number of curious cells; each is a 
hair-like filament, with a nuclear thickening either at its base 
or elsewhere along it, and these filaments stretch from one 
side right across the basal cavity towards the other (figs. 164, 
165). 

What the function of these extraordinary organs is I 
am quite unable to say. Lowne, searching for excretory 
organs in Calliphora, after denying the excretory function of 
the malpighian tubules, ascribed this function, without any 
definite reason, to the rectal glands. Hewitt observed them in 
the house-fly undergoing rhythmical contractions; in that 
insect they communicate with the body cavity, and he likewise 
concluded that they were excretory organs. The only fact in 
favour of this view, however, is their curious position; but 
the development of such organs in an insect already well pro- 
vided with excretory tubules, seems to contradict this view. 
In Vasoma the only communication with the body cavity that 
they may have is by means of the basal filamentous cells; 
but what their function is must remain, for the time, un- 
decided. In most insects two pairs of rectal glands seem to 
be present. 


The single salivary gland is composed of a relatively small 
number of thickened granular cells, with large granular 
nuclei. A few cells in the centre of the gland have a vacuo- 
lated or branched appearance, giving the interior of the gland 
a spongy structure. The gland lies in close contact with the 
great salivary duct, and the walls of this duct, in the neigh- 
bourhood of the gland, present the same vacuolated appear- 
ance as characterises the interior of the gland. The gland 


433 


secretion evidently enters the duct by percolating through this 
spongy tissue. 

In its other regions the duct is lined by flattened epithelial 
cells, which thicken near its opening. Here a number of 
muscles are attached, which are inserted at the other ends 
upon the walls of the head. 

The duct is lined by a chitin sheath, which presents 
spiral ridges similar to those seen in tracheae. 

The malpighian tubes (fig. 150) are eight thread-like 
cylindrical structures, each with a narrow flattened duct; the 
duct is formed essentially by the incomplete junction of 
embryonic cells in irregular pairs. Sometimes adjacent cells 
do not fuse completely and the canal in consequence extends 
between these also. 

The cytoplasm of the cells is very clear and homogeneous 
but often exhibits very large vacuoles. The nuclei are large 
and granular and have one or two medium-sized nucleoli. 


(2) The Intestine of the first Larval Insiar. 


In the larva there is a small mouth (fig. 1), the openings 
of the conical buccal cavity, which contains the minute sharp 
jaws (fig. 47). This leads behind into a long oesophagus 
(figs. 1, 140) which passes horizontally backwards through 
the circumoesophageal nerve ring, and opens in the third seg- 
ment into a great sac-like dilatation, the midgut (figs. 2, 140). 
The latter is endodermal, the oesophagus ectodermal in origin. 
The midgut occupies the greater part of the body extending 
backwards to the third last segment. Here it lies in close 
connection with the hindgut, but the two do not communicate 
till the time of defaecation, 7.e., till about one day before 
pupation. The midgut, indeed, is simply a blindly ending 
sac (fig. 143), and it is not till the last day of larval life that 
the unabsorbed food is discharged. 

From a portion of the oesophagus (foregut) the whole of 
the intestine of the imago as far back as the beginning of the 
stomach will develop during metamorphosis. From the great 
larval midgut the stomach of the imago is formed, while the 
small intestine and rectum are developed from the hindgut. 
The endodermal portion of the intestine is therefore much 
smaller in the adult than in the larval insect, and it is entirely 
from the ectoderm that the other structures—crop, gizzard, 
etc.—become developed. 


It is curious to observe how highly differentiated the 


intestine of the imago is, in an insect which rarely feeds, while 
the larva, which does nothing but feed, must content itself 
with so simple a structure. 7 
Opening into the base of the mouth is a median salivary 
duct, which soon divides into two parts. Each of these smaller 


434 


_ducts passes backwards and communicates with the two large 
salivary glands (fig. 142), which reach back to about the 
fourth body segment. The distal extremity is drawn out into 
a narrow prolongation of the main salivary gland. 

Running along the sides of the midgut, and opening into 
it posteriorly, is a pair of long moniliform tubes, the haepatic 
caeca. A third such tube, shorter, however, than these, 
opens into the midgut behind, and passes backwards over the 
rectum (fig. 140). The two lateral caeca have been observed 
by various investigators in the larvae of chalcid wasps. The 
third median caecum has not, so far as I am aware, hitherto 
been described. But the interpretation which is always placed 
on them is quite erroneous. They are regarded as malpighian 
tubes, but have in reality an entirely different function. For 
this interpretation see, for example, M. Havilarid (1920, 
1921). They are digestive glands, and in both structure (as 
will be seen later) and embryonic development, differ entirely 
from true malpighian tubes. The latter are ectodermal in 
origin; the structures that occur in the larva of Vasoma are 
outgrowths from the endodermal midgut. Furthermore, they 
have no opening to the exterior during larval life, and empty 
their secretions into the blindly-ending midgut. 

The necessity for such large haepatic caeca is clear when 
we consider the rapidity of feeding during larval life; the 
salivary glands are quite unable to cope with so great a 
quantity of food, and well-developed haepatic caeca are but 
to be expected. The secretion is poured into the posterior 
part. of the midgut, and the mixing with the engulfed food 
is the result of a remarkable forward peristalsis which can 
clearly be observed about once every fifteen to twenty seconds 
in the feeding larva. A peristaltic wave travels along, from 
behind, forwards, not only on the walls of the midgut, but 
also on the body surface itself, and this brings about a per- 
fect churning of the contents of the great food sac. 

When now we look for the true malpighian tubes, we 
fnd that they are absent, and that the larva_is entirely 
devoid of excretory organs. And unless excretion occurs by 
the diffusion of ammonia through the integument, no excretory 
_ activity goes on during active larval life. 

This fact, however, is less remarkable than it may at 
first sight appear to be. The larva is exceedingly sluggish, 
and is feeding upon food which has approximately the chemical 
composition of its own tissues. Practically the only energy 
expended is that which is needed in growth, and it is con- 
ceivable that the excretory products resulting from the neces- 
sary protein deaminisation accumulate in the blood during 
the five days of larval life. In the late larva numerous 


435 


crystals, evidently excretory in nature, begin to accumulate 
within the fat-body, and these do not disappear from there 
till the malpighian tubules are already well developed. Their 
disappearance coincides at that period with the appearance 
of undoubted urates in large quantity in the intestine. 

The loss of nitrogen as diffusible ammonia must, of 
course, not be disregarded, but the fact that actual excretory 
organs are absent cannot be doubted. 

It should be observed that this in no way supports the 
well-known statement of Lowne that the malpighian tubes of 


insects have a hepatic function. Urinary crystals often occur 


in immense numbers within the tubules of various insects, 
and their excretory function is established beyond doubt. 
Lowne, in searching for excretory organs, attributed this 
function to the rectal glands. He also regarded the periodic 
moulting as aiding in nitrogen excretion. The chitinous 
cuticle, however, which is shed is chemically an amino- 
polysaccharide, and contains less nitrogen than does protein ; 
if anything, its formation increases the proportion of nitrogen 
in the larva. | 


The histological differentiation of the various regions of 


‘of the larval intestine is not very marked in the first instar. 


The buccal cavity is lined by rather small clear cells, 64 
in thickness, each with a large clear nucleus containing a 
large karyosome but no nucleolus. Internally the buccal 
cavity is lined by a thick chitinous sheath (fig. 47). On the 
dorsal side of the buccal cavity are a number of circular 
muscles—large unicellular spindle-shaped structures inserted 
by their two ends upon the two lateral walls of the buccal 
cavity. In the first instar no distinct striations are visible, 
but in later larval life these differentiate. 


The oesophagus, which is lined by cuticle, is composed 
of a single layer of small cubical cells, presenting the usual 
undifferentiated appearance of the larval cells at this stage, 
viz., clear cytoplasm, and a vesicular nucleus with a large 
karyosome. The oesophagus projects slightly into the great 
midgut, and this serves as a valve to prevent any regurgi- 
tation of food during forward peristalsis. The oesophageal 
epithelium, just in front of the great midgut, is several cells . 
in thickness. The cells are slightly smaller than those found ~ 
elsewhere in the oesophagus, but are otherwise indistinguish- 
able from them. The slightly thickened ring which they form 
is the wmaginal disc of the oesophagus, from which the greater 
part of the foregut of the imago as far back as the stomach 
will develop during pupal life. 


436 


The midgut is lined by what appears to be a very delicate 
cuticle. The epithelial cells lining it are much larger than 
those occurring elsewhere in the intestine. The great accumu- 
lation of food within the midgut soon stretches them (fig. 10), 
and already at the end of the first instar they are becoming 
flattened; they measure at this stage about 68 in length 
(and breadth), lly in thickness. The cell cytoplasm is vacuo- 
lated and granular. The nucleus is large (17p) and is very 
heavily granular. 

At the base of many of these cells there is frequently 
to be observed a much smaller cell, spindle-shaped, about 
17p in length, 4u in height (fig. 10). Each has a large clear 
nucleus and a distinct karyosome, and represents an. un- 
differentiated non-functioning cell, which will become active 
during the defaecation period, and will form the great endo- 
dermal intestine of the early pupa. The anterior portion of 
this, as will be described later, will disintegrate, while the 
hinder will persist as the stomach of the imago. It is, there- 
fore, possible to speak of these cells when they occur in the 
posterior region of the intestine as imaginal Bs cells ; 
the more anterior ones cannot be thus described. TI shall 
refer to them later as replacing cells. 

External to the epithelium is a rough network of longi- 
tudinally and circularly disposed muscle fibres. 

The rectum (figs. 143, 185) is a prominent ectodermal 
ingrowth through the anus—a true proctodaeum, but does 
not yet open into the midgut. It is a rounded tube, lined 
by chitin, with a columnar, or, in places, cubical epithelium. 
Surrounding it is a layer of circularly disposed, as yet un- 
striated, muscle fibres. The epithelial and muscular cells still 
retain the usual undifferentiated appearance of clear cyto- 
plasm and large ‘“‘vesicular’’ nulcei. 

The cells at the anterior end of the proctodaeal invagina- 
tion fit tightly against the rear of the midgut, and form a 
layer several cells in thickness (fig. 143). The cells here are 
slightly smaller than elsewhere and constitute the zmaginal 
disc of the hindgut, from which the small intestine and rectum 
of the imago will later develop. 

The salivary glands. These consist each of a large sac- 
like structure—the secreting portion, and a duct of medium 
length, the two ducts uniting before opening into the mouth. 
In the first instar the common duct is rather flat and strap- 
like; along its middle passes a prominent canal, lined by a 
chitinous spiral intima, which is shed and reformed at each 
moult. Further behind, the duct becomes circular in section, 
and is composed of very small cells with the usual ‘‘first- 
instar’ appearance, viz., large clear ‘‘vesicular’’ nucleus, with 


437 
a big karyosome, and with clear hyaline cytoplasm. The 
glandular sac consists of about twelve very large, somewhat 
flattened cells, enclosing a large lumen. The cells are as 
much as 23» in diameter and have large granular nuclei 
lly in diameter. 

The hepatic caeca (fig. 10), which are usually falsely 
regarded as malpighian tubules, are composed of a number 
of very large rounded cells, arranged alternately in pairs; 
their union is such that a considerable space is left by the 
incomplete fusion of a cell with the one opposite it, while the 
fusion of cells with those behind them is always complete. 
The lumen is lined with myriads of minute cilia (figs. 10, 
146), whose movement drives the secretion backwards (or 
forwards, in the third caecum) into the intestine. The indi- 
vidual cells measure about 17% in diameter and have a 
remarkable resemblance to the cells of the fat-body. The 
nuclei are very large and heavily granular, and the cytoplasm 
faintly granular, and already at this early stage slightly 
vacuolated. - 

The hepatic caeca appear to be formed in the embryo as 
outgrowths from the midgut; this is indicated by the fact 
that in the first instar the third (posterior) caecum is present 
as a short, solid, robust projection from the rear of the mid- 
gut, and that it is only later that it acquires its ciliated 
lumen. ; 


(3) The Post-embryomec Development of the .Intestine. 


The feeding period of the larva (7.e., about the first three 
days of larval life) is characterized by the completion of 
differentiation of the cells of the first instar, by a great growth 
in cell size, and by a corresponding total absence of cell divi- 
sion, except in the case of those cells which constituted the 
“Imaginal tissues’? of the larval intestine, viz., (a) the 
oesophageal imaginal ring, surrounding the posterior part of 
the foregut; (6b) the small ‘‘replacing cells,’ as I shall 
designate them here, which lay scattered about at the bases 
of the large cells of the midgut; and (c) the imaginal disc 
at the anterior extremity of the rectum. 

The visible differentiation is not very marked; it con- 
cerns mostly the intestinal muscle cells which, though already 
functioning, have not yet adopted a striated appearance ; 


_ but before the end of the second instar this is always visible. 


(A) The Metamorphosis of the Foregut. 


During larval life there is a great increase in the size of 
the muscle and epithelial cells of the foregut; at each moult 


| the cuticle is shed and secreted anew. 


— 


438 


But shortly after feeding ceases the epithelial cells begin 
to degenerate. The nuclei are granular and greatly hyper- 
trophied, and possess a large nucleolus, so characteristic of 
the degenerating cells of Vasoma. In the defaecating larva 
the cytoplasm undergoes granular degeneration; or, at other 
times, it breaks up into larger globules, which, breaking from 
the cells, float about in the blood and are there engulfed 
by any leucocytes which happen to be present, or, if left to 
themselves, dissolve in the blood. 

These changes are accompanied by an active regeneration 
of the foregut (fig. 117). During larval life (though | can- 
not say definitely at which period of it) a proliferation of the 
cells of the oesophageal imaginal ring has occurred, and 
these, forsaking their ordinary cubical shape, elongate and 
become spindle-shaped (fig. 152). Continuing to divide mito- 
tically they bulge outwards, and at the same time extend 
forwards, and in the defaecating larva are to be seen actively 
replacing the disintegrating larval cells. Although I did 
not observe them penetrating these cells, as in the case of the 
myoblasts extending over dead muscles, yet it seems probable 
that they actively absorb the products of disintegration and 
grow at their expense, so near do they le to the dead larval 
cells. 

The oesophageal epithelium is partly regenerated also 
from another centre, v7z., the integumentary imaginal dises of 
the first segment; and in the defaecating larva these embryonic 
cells are to-be seen extending through the mouth inwards, 
between, or over the dead and disintegrating cells of the larval 
epithelium, while the proliferating cells of the oesophageal 
ring extend forwards to meet them (fig. 117). About four 
hours after defaecation the two have met. 

Meanwhile there has begun a proliferation of certain 
myoblast cells, which le during the whole larval life scat- 
tered about in the head in the neighbourhood of the mouth; 
these extend backwards as a loose column of very long 
spindle-shaped cells, drawn out in long thread-like processes 
at either end (fig. 117). They form the musculature of the 
anterior part of the oesophagus. Others are to be seen behind 
the oesophagus; the muscles which they form are differently 
disposed from those of the anterior side of the oesophagus, and 
will be considered later. The development of the great 
pharyngeal dilator muscles is considered in connection with 

_the development of the general muscular system. 


During the remainder of larval life the cells, occasionally 


still dividing, settle down, and growing in size, co-operate to 
form a single epithelium—that of the adult oesophagus. This 
development is accompanied by a great bending downwards 


439 


and backwards of the head, as already described, and the 
result is a total change in the course taken by the oesophagus. 
It now passes not directly backwards, but first upwards and 
forwards actually, and only then begins gradually to bend 
backwards. Meanwhile the cells of the circumoesophageal 
imaginal ring have continued to proliferate, and in the fresh 
pupa form a great cone of cells, attached behind to the 
anterior end of the foregut, and ending, in front, just behind 
the brain (fig. 154). The structure is composed of two layers 
—an inner of long columnar cells, all tightly compressed and 
arranged radially around a very narrow central lumen. Out- 
side this is a second layer one or more cells in thickness, 
the individual cells are much smaller here (fig. 153). From 
the great inner layer the succeeding portion of the intestine 
as far back as the gizzard is soon to develop. 

The smaller outer layer has a much humbler future ; when 
the cells of the great inner layer have migrated backwards 
(a process which commences several hours after pupation), 
the outer layer cells extend round to the lower side of the 
oesophagus and form a rather thick column there in front of 
the neck, but not extending as far downwards as the brain. 

Jt is in connection with this structure that the developing 
myoblasts of the rear of the oesophagus. now come; these 
cells, having united end to end during late larval life, now 
form several rows of cells, inserted behind all upon the sub- 
oesophageal cell column, and in front at various points on 
the rear of the oesophagus. Adjacent cell walls now break 
down, and each row forms a single multinucleated syncytium. 
During the third day of pupal life these syncytia develop 
spiral striations and form the post-oesophageal muscles of the 


-imago. Meanwhile, the sub-oesophageal cell column, upon 


which the columns of myoblasts are all inserted, begins, in 
the thirty-six hour pupa, to chitinise internally. Chitinisation 
continues and the chitin rod fuses with two other very short 
processes which have grown out from the rear of the head, 
close to the neck. By this means, the musculature of the rear 
of the oesophagus obtains a very firm support. 

Already in the fresh pupa, the myoblasts of the anterior 
part of the oesophagus have disposed themselves longitudinally 
or circularly in the position they are to occupy in the imago; 
several cells usually fuse to form small syncytia, and, under- 


_ going the usual differentiation, form the striated muscles of 
the adult oesophagus. 


It remains only to note that these changes are accom- 


_ panied by a new secretion of chitinous cuticle within the 
lumen of the oesophagus, the old having been drawn out 
_ through the mouth at the pupal moult. The development of 


440 


the rest of the foregut does not occur till a series of remark- 
able processes have taken place in the anterior half of the 
midgut; only then does the foregut extend backwards, and, 
occupying the place of the anterior half of the true (endo- 
dermal) midguts of the larva and early pupa, fuses with the 
posterior half, which remains as the stomach, being all that 
survives of the old midgut. The midgut will, therefore, most 
conveniently be considered first. 


(B) The Metamorphosis of the Midgut and the Development 
of the Post-oesophayeal part of the Foregut. 


I have already described the midgut as composed of a 
single layer of large flattened cells, with smaller cells at their 
' bases. As many of these smaller ‘cells do not survive in the 
imago, I shall refer to them here as replacing cells. 

Neither larval nor replacing cells, so far as I can observe, 
itt proliferate during larval life. The Jarval cells grow enormously 
in size, and, through the pressure exerted upon them by the 
contents of the midgut, are seen, at the end of the third day, 
as great flat cells, with smaller replacing cells at their bases. 
_ But during the third day these replacing cells begin slowly 

to divide by (mitosis) while the great larval cells remain 
inactive. 

But at the end of the fourth day of larval life a great 
change begins. At about this time the rectal ingrowth at 
last fuses with the midgut; and this event is marked by the 
commencement of a series of contractions of the muscles of 
the intestine which gradually drives the undigested food, 
which has accumulated here during larval life, to the exterior. 
This is the defaecation period of the larva, and lasts from 
one to two hours. But it is apparently under the pressure — 
exerted by the muscles that a remarkable process.of-.dis- 
integration of the epithelium of the larval midgut begins. 
The disappearance of the faecal material allows the cells to 
return to the cubical conditions in which they existed in the 
new-born larva. 

In the cytoplasm of the midgut epithelial cells before 
defaecation, vacuoles were already becoming numerous; 
usually, however, it was quite granular and showed obvious 
signs of degeneration. Not only had the cells grown greatly, 
but the nuclei had greatly hypertrophied, and were to be 
seen as long, irregular, faintly granular, chromatin masses, 
devoid of nucleoli. When the cells contract these long nuclei 
become bent; and a section presents the curious but false 
_ appearance of large multinucleated cells. 

But at defaecation the vacuolation becomes very much 
more marked (fig. 144); the vacuoles consist perhaps of fatty 


44] 


material and occasionally contain feebly staining grains. 
They are suspended in a very faintly granular ‘‘spongioplasm.”’ 
As the pressure exerted by the muscles increases the cells 
begin to project irregularly into the intestinal lumen; and 
then a very remarkable thing is to be observed. The spongio- 
plasm, together at times with the vacuoles, begins to ooze out 
through the cell membrane, and hangs as one or more large 
drops, suspended in the intestine from the degenerating cells 
(fig. 144). The process commences in the anterior part of the 
midgut, but soon extends right along it, as the faecal con- 
tents are gradually voided. The products of degeneration are 
themselves, however, retained in the lumen of the intestine. 
There they granulate and are seen sometimes as little balls 
of grains, at other times as a fine dust. Eventually the whole 
of the cytoplasm’ is cast into the now very contracted lumen 
of the midgut; and all that remains is the cell membrane 
containing a very degenerate looking clump of chromatin 
grains. But these are soon added to the mass of débris which 
now consists of fine grains, of small clusters of grains, of 
fragments of nuclei, and of the contracted walls of the dead 
cells; all forming a dark mass that now occupies the lumen 
of the intestine. In the larva eight hours after defaecation 
these changes are complete, and all the old larval epithelium 
has disappeared, with the exception of a narrow strip of 
dead cells running along either side of the intestine from one 
end to the other (fig. 146). The temporary retention of these 
cells is an extraordinary adaptation for bringing about the 
destruction of the hepatic caeca; their fate will be described 
later. 


It is necessary to return now to the replacing cells. In 
_ the defaecating larva these cells have begun to proliferate by 
mitosis, and by the time the larval cells have lost most of 
their cytoplasm (1.€., about four hours after defaecation), 
these have formed a completely new epithelium, closely sur- 
rounding the degenerate larval epithelium. This gives the 
intestine the false appearance of having a functional two- 
layered epithelium (fig. 145). But as the larval epithelium 
disappears more and more, the cells of the new epithelium 
increase in size, and in the eight-hour pupa alone persist, 
except for the two thick bands of dead cells on either side 
of the intestine, close beside the great hepatic caeca. 


But shortly after this an extraordinary thing is to be 
observed. The cells of the renovated epithelium, growing 
in size, begin to push the two longitudinal columns of dead 
cells into the lumen of the intestine. To each of these columns 
—the sole remains of the epithelium of the larval midgut— 
the hepatic caeca, which have now grown 70y in thickness, 

a | 


442 


with great granular nuclei 134 in diameter, are connected 
along their length by means of the fine membranous peri- 
toneum (fig. 146). And as the growing epithelium forces these 
remains of the old intestine into the lumen, the hepatic caeca 
are pulled bodily in with them along the whole.lensth of 
the intestine. In the larva sixteen hours after defaecation 
the hepatic caeca are being slowly but surely( engulfed (figs. 
147, 148), and six hours later have entirely vanished. To 
the débris within the alimentary canal is also added the third 
(posterior) caecum. This becomes drawn into the midgut in 
a very similar manner; dead larval epithelial cells at its base 
fail to disintegrate, and the surrounding cells of the new 
epithelium pushing these cells inwards cause the third caecum 
to be slowly drawn into the lumen where it disintegrates 
along with the other disorganized tissues. 


Even before being absorbed the hepatic caeca show signs 
of degeneration, small globules of cytoplasm being thrown 
into their lumen. 

The function of the renovated epithelium appears to 
be to absorb this débris (fig. 154), perhaps after it has been 
digested by the enzymes liberated from the disintegrated 
hepatic caeca. At any rate, a marked absorption of the 
débris commences in the fresh pupa. Muscular contractions 
in this region later drive part of the contents into the posterior 
portion of the gut, and here the apparently indigestible 
cell membranes of the old larval epithelium accumulate, and 
may persist in small quantity till the emergence of the wasp, 
when they are voided through the anus. But the greater 
part of the fine granular débris soon disappears. 

Meanwhile in the late hours of larval life a few cells 
from the rear of the great conical circumoesophageal imaginal 
ring have grown in as a short solid mass of cells into the 
anterior end of the midgut (figs. 153, 154). They push the 
adjacent intestinal cells along with them and the two co- 
operate to form a temporary obstruction which prevents the 
débris within the intestine (especially when the muscles of 
the anterior portion contract) from entering the foregut. This 

‘plug’? does not, however, develop till shortly before pupa- 
tion, and closure “of the anterior region of the midgut during 
the period which intervenes between defaecation and this, is 
brought about by a muscular contraction here, which causes 
considerable folding of the epithelium, and a consequent 
closure of the passage. The general nature of the regenerated 
midgut is seen in fig. 154. 

The cells of the newly formed epithelium, which have 
now attained quite a large size, having performed their func- 
tion, now begin to disintegrate, oe as a second time an 


Ao TT en 


ba 


a Se 


443 


extensive destruction of the midgut occurs. The process com- 
mences shortly after pupation, but is quite different from what 
occurred in the larval midgut. The whole epithelium of the 
anterior half of the midgut begins to break up into a mass 
of rough granules. The muscle fibres of the intestine, losing 
their striations, join in the general process of disintegration, 
and what was a few hours earlier an actively functioning 


tissue, is now a loose accumulation of granular débris. This_ 


time, however, the leucocytes act, and_swarming into the 
disintegrated mass rapidly absorb it (fig. 153)); after six hours 
not a trace of the temporary midgut remains. The small 
conical projection of the foregut takes part in the general 
destruction, and nothing of the renovated midgut remains 
except the posterior half, in which no disintegration whatever 
has occurred. This portion remains with but little change 
as the stomach of the adult insect. Its structure has been 
referred to already. 

But before the anterior half of the midgut has had time 
to disappear entirely, the cells of the inner layer of the great 


conical circumoesophageal imaginal ring have sprung into 


activity; they grow rapidly, moving evidently by amoeboid 
action, along the pathway afforded by the disintegrating 
mass, and, extending right through the thorax and anterior 
abdominal segments, at last reach, in the eight-hour pupa, 
the .anterior end of the hinder half of the midgut which has 
survived these violent scenes unchanged. From this newly 
formed structure the post-oesophageal part of the foregut 
soon begins to differentiate. 

In the thoracic and propodeal regions it is formed, and 
persists, as a very fine, almost capillary, tube, 8u to 10y in 
diameter (cf. fig. 156). The cells, which are at first irregular 
and embryonic in appearance, soon elongate, grow spindle- 
shaped, and dispose themselves longitudinally. They seem 
to lose part of their cytoplasm later in pupal life, and in the 
adult insect appear almost devoid of it. 

But the abdominal portion of the foregut undergoes a 
much more complex differentiation. In the eight-hours pupa 
its posterior extremity is seen as a thick-walled, somewhat 
conical and slightly dilated chamber (fig. 155). This will 
develop into the crop, the gizzard, and the ‘‘drum-shaped’”’ 
chamber. Its epithelium is composed of long columnar cells, 
which gradually merge, in the region of the petiole, into 
those of the narrow capillary portion. Surrounding the 
structure is already to be seen a number of cells forming a 
distinct layer of myoblasts. The lumen does not yet com- 
municate with that of the stomach, but ends blindly. The 
hinder part of this lumen is rather constricted and will 


=. < 


444 


form the cavity of the gizzard and ‘‘drum-shaped’’ chamber. 
The more anterior part 1s more widely dilated. Here the 
crop will develop. 


The development of the crop is very curious. The epi- 
thelial cells entirely lose their columnar character; they 
flatten out more and more, and in the pupa one day old have 
become highly wrinkled. The flattening continues, and 
instead of a small conical chamber with very thick walls 
there is formed a highly distensible collapsed bag with very 
thin paper-like walls (fig. 156). The cell differentiation 
closely resembles that observed in the cells of the differ- 
entiating wing epithelium. 

The gizzard rapidly differentiates. Already at the end 
of the first day its lumen has become triradiate; this is 
brought about by the epithelium arranging itself in the form 
of three short longitudinal plates; bent along their longi- 
tudinal midline. The epithelial cells are still embryonic in 
appearance. The myoblasts have already arranged them- 
selves in their definite positions. Their future development is 
exactly the same as occurs in the case of other muscles and 
need be referred to no more. 


In the thirty-six hour pupa the three bent epithelial plates 
are beginning to secrete chitin on their inner walls; and the 
posterior portion, where chitinisation does not occur, is 
observed to be marked off as a small rounded chamber, into 
which a short ‘‘filament’’ projects from the hinder wall 
(fig. 156). 

During the next day the chitinisation strengthens, and 
with the appearance of the muscle striations the gizzard attains 
its adult proportions. 


Cell proliferation of the drum-shaped chamber occurs 
also at this time, and at last a communication between foregut 
and stomach is established (fig. 157). 


(C) The Metamorphosis of the Hindgut. 


The walls of the rectum undergo the same changes during 
larval life as occur elsewhere in the larva, 7.e., there is a 
growth in cell size, in the absence of cell division; the muscle 
fibres during the second instar gradually acquire striation. 

But about half a day after feeding ceases, the cells of the 
anterior end of the rectum, which constitute the imaginal disc, 
become active, and proliferating greatly at last bring about a 
junction of the cavities of the midgut and hindgut. The actual 
opening is large and funnel-shaped. Not till this time, there- 
fore, does the embryonic proctodaeal invagination open into 


—— 


445 


the archenteron ; usually, in other insects, this occurs during 
embryonic life. | 

This event is quickly followed by muscular contractions in 
the midgut, and two hours later the whole of the faecal 
matter, which has accumulated during the three days of 
active feeding, is voided. The rectal musculature takes part 
in the process; only a small part of the faecal matter at a 
time is passed into the rectum, and this, under the pressure 
of the muscular walls, is rounded off into a little pellet, which 
is forced slowly along the rectum. ! 

Meanwhile the epithelial cells have begun to degenerate. 
The nuclei are large and granular, the karyosomes having 
scattered their material through the nucleoplasm as this 
gradually hypertrophied. The cytoplasm then undergoes 
granular degeneration. In the defaecating larva these 
granules cluster together in little balls and breaking through 


the cell membrane float about in the blood stream, where they 


may become engulfed by phagocytes; or, if these are not 
present at the time, simply dissolve in the blood plasma. At 


’ other times the cytoplasm becomes broken up into a number 


of larger hyaline globules, like those of the integumental cells. 
They share the same fate as do the balls of granules. 

Meanwhile, the cells of the rectal imaginal ring grow 
backwards as well as forwards, and as the rectal epithelium 
disintegrates they rapidly replace it. Already in the defae- 
cating larva the epithelium of the anterior quarter of the 
rectum is composed entirely of embryonic cells, and these, 
dividing mitotically, are actively growing backwards, replacing 
the epithelial cells as these disintegrate more and more. A 
few hours later the whole larval epithelium has disappeared, - 
and a loose layer of spindle-shaped embryonic cells has re- 
placed it. The muscle layer does not disappear till a few | 
hours after pupation. ! 

The cells of the renovated epithelium now consolidate 
their position. In the larva some eight hours before pupa- 
tion a cuticle is being secreted between the old cuticle and the 
epithelium, and when the larva moults the old cuticle of the 
last larval instar is drawn out through the anus. 

In the fresh pupa myoblast cells, which were present in 
only small numbers during larval life, having proliferated 
considerably during the last twenty-four hours, now form a 
distinct layer outside the rectum, and it is not till several 
hours later that the old larval muscles begin to disintegrate 


and. become phagocytised. 

The renovated arr epithelium now begins to differ- 
entiate into two regions; the small spacious rectum behind 
and the small intestine in front. : 


446 


The cells in the mid-region of the hindgut are, in the 
early pupa, in a state of rapid proliferation; and this, con- 
tinuing into the next day, produces a considerable bending of | 
the anterior region. That portion behind the centre of pro- | 
liferation is the rectum; the portion anterior to it, the small 
intestine. The amount of this proliferation seems to vary 
considerably, so that, while the intestine is sometimes quite 
bent upon itself, at other times the bending is far less marked. 

Having peer at its maximum size, the cells of the 
small intestine begin to differentiate; but the differentiation 
never becomes very marked, and the epithelium remains as 
a single layer of elongated loosely arranged cells, on whose 
surface an equally irregular chitin sheath is secreted. The 
organ is evidently capable of considerable stretching. 


Already in the fresh pupa the rectal region of the hind- 
gut is distinguishable from the small intestine by the ‘‘spindle 
shape’ of its epithelial cells. The tube is already more dis- 
tended than the anterior portion, but six hours later attains 

(its adult proportions. It is not till six to eight hours after 
pupation that the larval muscles finally disappear by phago- 
_cytosis, after having undergone globular degeneration. 

The adult muscles develop in the usual manner. 


The Rectal Glands. 


During the last hours of larval life the cells of the 
anterior region of the rectal portion of the hindgut begin to 
proliferate and grow in the -form of two small clumps of 
elongated cells into the cavity of the enlarging rectum (fig. 
161). These are the rudiments of the single pair of rectal 
glands, and have already attained to a considerable size in 
the fresh pupa. 

The rectal glands grow considerably in size. The elongated 
cells dispose themselves in a single layer with their long axes 
vertical to the surface of the gland; the whole structure has, 
in the eight-hour pupa, a short cylindrical shape. It is solid 
except below, where there is a cavity lying loosely in which 
is a large number of much smaller cells. They are the 
elongated filamentous cells already described. A cuticle is in 
process of secretion. The rectal gland lies in close contact 
with the wall of the rectum from which it has been developed, 
and there appears to be no communication through it, 
between the cavity of the rectal gland and the haemocoele. 

During the rest of the first day the rectal gland elongates 
considerably, by growth of its cells, not by their proliferation. 
The small basal cells now dispose themselves in a ring at the 
base of the gland; in the twenty-one hour pupa some have 


447 


developed their remarkable filamentous structure; others are 
in process of dividing. This leaves the cavity, in which they 
lay, devoid of cells, and at this stage a canal, formed by the 
incomplete fusion of the bases of the large cells, is developed 
right along the axis of the gland, and opens below into the 
large basal chamber. The great elongated cells have mean- 
while begun to fuse on their outer surface, and from now 
on the rectal gland may be divided into an outer syncytial 
cortical portion, surrounding an inner medullary region in | 
which cell walls are still well marked. The cortical portion 
is formed by the fusion of the outer ends of the elongated 
cells; the medulla is the region which becomes differentiated 
by the failure of the cells to fuse here. 

In the thirty-six hour pupa the glands have grown to their 

maximum size. Cortex and medulla—are—very~ clearly seen 
(fig. 163). The central canal is prominent; the basal cells 
have differentiated into their adult filamentous condition. 
The cells of the rectum at the base of the two glands have 
proliferated a little to form two thickened pads; a trachea 
soon penetrates the rectal wall here. During the third day, 
the thickened pad begins to chitinise. 
Even in the fifty-six hour pupa the medullary region is 
distinguishable, but from now on the syncytium becomes more 
and more developed, and in the mature organ no distinction 
can usually be drawn between medulla and cortex. 

In the advanced pupa the large elongated cells begin to 
develop nucleoli of extraordinary dimensions (fig. 166). Some- 
times they occupy nearly the whole nuclear space. 


The Malmghian Tubes. | 


In the adult larva shortly after the cessation of feeding, 
the malpighian tubes become visible as small papillae on the 
anterior end of the hindgut (fig. 160). Here the cells of the 
imaginal ring have begun to develop, and it is not till this 
time that definite malpighian tubes can be observed. 

The papillae begin to grow with extraordinary rapidity, 
and by the time the larva defeacates (1.e., twelve hours later) 
they are visible as long thin threads (fig. 151), sometimes 
reaching almost to the dorsal body surface. From the first 
they have a narrow lumen. The walls are one cell in thick- 
ness; the cells are roughly cubical and fit loosely together. 
The tubes measure about 6u in diameter. 

The tubes continue to grow in length, but not in thick- 
ness. In the fresh pupa the cells, which had previously the 
usual embryonic features—large karyosome, ‘‘vesicular’’ 
nuclei, hyaline cytoplasm—now begin to show signs of dif- 
ferentiation. The nuclei become granular, and the cytoplasm 


448 


becomes uniformly slightly vacuolated. Nucleoli are not yet 
present. 

The tubes from now on grow mainly in thickness, and in 
the thirty-six hour pupa have attained their adult propor- 
tions. With the exception of an absence of nucleoli the cells 
are, to all visible appearances, in their adult condition. 


The appearance of the malpighian tubes is followed 
shortly by the deposit of excretory material within the 
stomach (midgut). During larval life, as I have pointed out 
above, no removal of excretory substances appears to occur; 
towards the end of larval life, when the processes of growth 
necessitate a considerable deaminisation of the proteins of 
the disintegrating tissues, and perhaps of protein reserves 
within the fat-body, crystals, which are regarded as urates, 
accumulate in the fat-body and nucleoli of various tissue 
cells. On the other hand, a microscopic examination of the 
contents of the midgut of the larva shows no trace of these. 
But after the first day of pupal life small crystals begin to 
appear in the stomach; in the thirty-six hour pupa they 
increase in number and size; and from now on the stomach 
becomes a depositing place for the excreted urates and the 
undigested hulks of the old larval epithelial midgut cells. 

In many instances (¢.g., the silkworm) the urates are to be 
observed within the malpighian tubes as minute crystals. In 
Nasonia, however, they do not crystallize out till reaching 
the stomach. Here some of the crystals actually are far 
wider than the lumina of the tubules, and have grown in size 
within the stomach. 

At the time these crystals begin to appear in the intestine 
those of the fat-body and nucleoli disappear, and though the 
crystals in the two places have no resemblance to one 
another, yet it is probable that the two events are closely 
related. Pérez (1920) observed that the “‘pseudonuclei’’ of 
the large storage granules (“‘albuminoid grains’’) disappeared 
when the urates began to accumulate in the rectum of 
metamorphosing insects; and came to the same conclusion as 
that expressed above, basing his view on the experiments. of 


Marchal, who was able to convert these ‘‘pseudonuclei’’ into 


crystals by treatment with acid. 

Finally, when the wasp hatches, these excretory crystals 
and any other contents of the stomach are thrown out. 

Crystals similar to these form the creamy or pink material 
excreted by insects shortly after emergence. They are especi- 
ally well seen in the silkworm, where they can be gathered 
in considerable quantities. They give the murexide test for 
uric acid; a faint ammonia reaction can also be obtained with 
Nessler’s solution. 


: 
| 


449 | 
The Salivary Glands. 

The main changes undergone by the salivary glands 
during larval life are a great growth in the size of the con- 
stituent cells. In the adult larva they are as much as 57 
in length, 284 in breadth; they are highly vacuolated and 
have gigantic granular nuclei 30u long, 17» broad, and usually 
contain several small nucleoli. 

The duct cells also grow largely in size; their spiral 
intima is shed and reformed at each larval moult. 

While the glands do not disappear till early in the pupal 


_ period, the median duct is actively disintegrating already in 


the defaecating larva. The larval cells have met the same 


fate as those of the oesophagus, 7.e., they have degenerated 
and the products of degeneration have been cast, in part, at 
any rate, into the blood stream. Renovation of the median 
duct quickly ensues (fig. 117); embryonic cells growing in- 
wards and downwards from the regenerating epithelium of 
the mouth and pharynx pass among the disintegrating cells, 
nourishing themselves perhaps; in part, at their expense. 

~The cells, however, do not grow back along the duct 
beyond its point of bifurcation ; forsaking the old larval 
salivary duct here (the larval ducts at this point do not 
appear to be dead yet) they grow as a slender, hollow column 
up the back of the head and terminate in the neck. From 
the duct at a point about one-quarter its length from the 
mouth, the salivary gland of the imago develops in the second 
day of pupal life. I have not observed the process, ebut it , 
seems unlikely that the gland should be formed in any other 
way than by a thickening of the duct. 

Meanwhile, the remainder of the larval salivary glands 
disappear. In the larva shortly before pupation the 
greatly hypertrophied cells are to be observed undergoing 
obvious degeneration. Numerous minute globules are to be 
seen oozing out from the gland cells (fig. 149) into the cavity 
of the gland, in the same manner as I have described above 
in the hepatic caeca. The cytoplasm is even more highly 
vacuolated than usual. But about six hours later (four-hour 


_ pupa) the cells have entered into a state of granular dis- 
_ integration. Parts of the cells are in a condition of fine 


débris; other parts seem to have till now maintained their 
structure. The whole organ is very fragile, and I have 
observed a case in which the tracheae of the forewings, grow- 
ing downwards from the main trunks, have torn off a portion 
of the disintegrating tissue and carried it along with them 
(fig. 88). 

Lying within the e disintegrating salivary glands are great 

bers ytes,” “actively engaged 1 in clearing away the 


_ 450 


débris (fig. 88). Indeed, it is difficult to state how much 
_of the disintegrated tissue is removed in this manner, and 
how much in the more direct way of solution in the blood 
stream. 

Of the occurrence of phagocytosis of the gland cells, how- 
ever, there can be no doubt whatever. Indeed, the salivary 
glands, in their degeneration, offer as clear an example of 
phagocytosis as it is possible to wish for. But their death, 
degeneration, and even partial disintegration previous to 
phagocytosis are equally clear. 

The bifurcated portion of the salivary ducts disintegrates 
at about the same time. The leucocytes, having removed 
the débris from the glands, now move forwards and absorb 
the ducts also, and in the pupa six hours old no trace. of the 
larval glands is any longer to be recognized. 

The metamorphosis of the insect intestine has been the 
subject of a number of distinct investigations, to which I 
can refer but briefly here. Weismann, Kowalevsky, Lowne, 
and more recently Pérez (1910), have carefully examined the 
process in Calliphora, and it seems to differ but little from 
that of Vasoma, so far as essential characters are concerned. 

Deegener has investigated the metamorphosis of the 
intestine in the Coleoptera Hydrophilus (1910), Cybister 
(1904), and in Malacosoma (1908); while Rengel has made 
observations on Tenebrio molitor, and several water-beetles. 

The metamorphosis of the intestine of the silkworm has 
been investigated by Verson (1898, 1905); Poyarkoff (1910) 
examined that of a Chrysomelid beetle Galeruca; and Russ 
(1908) studied it in the Trichoptera. 

The observations of these workers differ considerably, 
and while differences in the material dealt with may account 
in part for the discrepancies, misrepresentations must not be 
forgotten. Thus, while the formation of a replacing epithelium 
in the midgut, confined to the pupal period, and similar to 
that occurring in Wasoma, is fairly frequent, it appears to 
be absent in some forms. For example, Deegener could not 
observe it in Malacosoma; in the Trichoptera Russ failed to 
observe it, and regarded a constricted part of the imaginal 


midgut as functioning in its place. Verson was not able to ~ 


see it in the silkworm; since its function is to absorb the 
products of degeneration of the larval midgut, its absence in 
the silkworm may be correlated with the voiding, as observed 
by Verson, of these degeneration products through the anus, 
shortly before pupation. 

If the observations of Verson are correct, the process offers 
a curious type of inefficiency—the waste of certain useful 
storage materials—which does not occur in Nasoma. In 


451 


Cybister Deegener describes a second temporary epithelium, 
found late in larval life, and inserted between the old larval 
and temporary pupal epithelia. It should be noted that the 
temporary pupal epithelium is a very transient structure, and 
that its absence in some insects is only apparent. 

In Galeruca Poyarkoff described a very interesting 
rejuvenation in the cells of the fore- and hindguts, similar 
to that occurring in the integument. In the silkworm, 
according to Verson, and also probably in the Coleoptera, 
there is a cell proliferation in the epithelium of the fore- and 
hindguts previous to each larval moult. This is evidently 
comparable with the proliferation of the fat cells of Galeruca 
as observed by Poyarkoff. Its significance will be explained 
in the second portion of this paper. 


The malpighian tubes are of special interest. In Vasoma, ais 


and_evidently in many other chalcid wasps, they are absent 
in the larva. In Calliphora Lowne believed them to undergo 
phagocytic destruction ; but more recently Pérez has observed 


them to undergo during pupation a remarkable process of 


dedifferentiation, followed later by redifferentiation into _ 


imaginal organs, In Galeruca, however, there is.a Bae: 


of Seen a to form the aaaeinal organs. 


THE DUCTLESS GLANDS. 


Under this heading three structures will be considered :— 
(a) The oenocytes; (b) a pair of lateral intestinal glands, 
occurring only in the larva, and which have not, I believe, 
hitherto been observed; and (c) a pair of dorsal abdominal 
glands, functional only in the adult insect, and also, so far 
as I am aware, hitherto unrecorded. 


The Oenocytes. 

The term oenocyte was first applied by Wielowiejsky in 
1886 to certain very large cells scattered about among the 
cells of the fat-body of Corethra. Tichomiroff had already 
noticed then in 1882 in the silkworm; he observed their 
proximity to the tracheae and regarded them as of ectodermal 
origin. Wheeler in 1892 found them to delaminate from the 
ectoderm; Weissenberg in 1907, studying them in a chalcid 
wasp Torymus, came to a similar conclusion. Finally, Nelson 
(1915), examining them in the embryo of the honey bee, saw 
them invaginating in close relation with the stigmatic trunks 
from the lateral ectoderm. Berlese has described them in 
the hymenopteran Tapinoma as a pair of small masses of 
cells in the fifth to the eleventh segments. 


i 
§ 


452 


Their segmental nature, together with their peculiar mode 
of development, strongly support the view of Lowne (1890) 
that they are homologous with the néphridia of annulata: 

Their function is not definitely known; Berlese regarded 
them as excretory organs. Glaser, in 1912, extracted an 
oxidising enzyme from them. They appear to be scattered, 
as a rule, fairly uniformly amongst the cells of the fat-body, 
and this suggests that they are in some way related function- 
ally to this structure. Perhaps their secretion contains some 
enzymes which drive the storage substances of the fat cells 
into solution, when the organism needs them. 

Their behaviour during development seems to vary with 
different insects. In Calliphora, Lowne (1890) observed their 
histolysis during the early pupal period. Pérez (1910) 
observed their phagocytosis and described the oenocytes of 
the adult fly as arising from certain smaller imaginal 
oenocytes, present in the body cavity. 

In Galeruca, Poyarkoff (1910) described certain larval 
oenocytes as undergoing phagocytic destruction at the end of 
larval life; others bud off numerous daughter cells, which 
become the oenocytes of the adult insect; the remaining por- 
tion of such a larval oenocyte becomes, after budding, the 
victim of the phagocytic activity of the fat cells and 
leucocytes. 

In the ant Formica rufa, Pérez (1902) described a some- 
what similar budding at the end of larval life. But in Call- 
phora this does not occur. In the honey bee, according to 
the observations of Nelson (1915), no cell division takes place 
in the oenocytes, once they have left the ectoderm from which 
they were formed. Similarly in the chalcid wasp Torymus, 
Weissenberg (1907) observed phagocytosis of the larval 
oenocytes at the end of larval life, while the cells of the 
adult wasp were produced from certain ‘‘imaginal oenocytes” 
lying within the body cavity. 


When the mature larva of Wasoma is examined the 
oenocytes are seen as about eight to twelve large cells in 
each segment from the third to the twelfth, lying on either 
side of the intestine, and singly distributed, fairly evenly, 
through the fat-body. They are the oenocytes which have 


_ functioned during larval life, and are present already in the 


newly hatched larva. They are not unlike the cells of the 
fat-body in appearance at first, but as the latter accumulate 
storage products the resemblance soon disappears. Towards 
the end of the first instar the cells have grown a little; they 
are apparently spherical and measure 12 to 13y in diameter. 


453 


The cytoplasm is fairly clear, though with a very faint indica- 
tion of granulation; the outermost ee are faintly vacuo- 
lated. The nucleus is large, measuring 54 to 6p in diameter ; 
its chromatic contents are fairly evenly scattered and there 
is one large central karyosome. 

During larval life the oenocytes grow considerably in 
size, having in the mature larva a diameter of about 45u 
(sometimes as much as 55u). The nucleus grows in propor- 
tion ; its diameter is about 25u. There is no evidence, there- 
fore, of any marked difference in the nucleo-cytoplasmic ratio 
in the young and old larvae, so far, at any rate, as the actual 
volumes of the two materials are concerned. Whether there 
has been an increase in the quantity of chromatic material is 
more difficult to observe; there seems, however, to be no 
evidence that such has occurred; the great karyosome has 
disappeared and scattered its contents throughout the 
. enlarged nuclear space; in its place, however, are to be seen 
one or a few prominent nucleoli often containing crystals; 
sometimes as many as twenty smaller ones are present 
instead. The cytoplasm is generally faintly granular, and 
usually heavily vacuolated in its outer regions (fig. 76). 
Generally the oenocytes are spherical, but often they become 
partly indented by other organs—tracheae or muscles—against 
which they have been pressed as they gradually grew in size. 

In the late larva and in the earliest hours of the pupa 
these cells degenerate and finally disappear, and all stages of 
degeneration may be observed during this period. Often in 
the mature larva the oenocytes may show a division of the 
cytoplasm into an inner heavily granular and an outer less 
granular zone, which is to be looked upon, apparently, as the 
beginning of disorganization. But it is not till the time of 
pupation that actual disintegration occurs. 

Usually the surrounding fat cells prevent the approach 
of leucocytes, and the oenocytes disintegrate spontaneously , 
large rents appear in the cytoplasm, and these develop into 
great holes; and at other times the’ whole cytoplasm 
degenerates into a fine powder, which is cast into the blood 
tfips.) 1765" 177). 

But when the surrounding fat cells are not so densely 
packed as to prevent the leucocytes from taking part in the 
process, the latter appear (fig. 178), and, before chemical 
disintegration has had time to occur, they overwhelm the 
cells and, eating large pieces out of their substance, eventu- 
ally ue them. So far as I could observe, the larval 
oenocytes do not persist beyond the early hours of the pupal 
stage. 

Meanwhile, the oenocytes of the adult wasp have been 
developing. They are represented, in the defaecating larva, 


454 


by small groups of closely packed, rather large cells (fig. 92) ; 
they still lie quite close to the integument, and though they 
occur elsewhere, are best seen at the posterior end of the larva. 
They do not occur in the head. 

In the larva of the first instar they can be seen (fig. 175) 
as a small cluster of rounded cells, which have just grown 
down into the body cavity from the ectoderm of the integu- 
ment. The cells measure about 104 in diameter, the nuclei 
5hu. They have a large karyosome, but the chromatin is not 
markedly scattered through the nucleus. 

During larval life they grow downwards and increase 
in size, attaining in the defaecating larva a diameter of 15n, 
while the nuclear diameter has increased to 8u; a small 
(plastin) nucleolus has begun to appear. The cytoplasm is 
homogeneous. I could not observe any increase in the wwmber 
of the oenocytes, however. 


But as the larval oenocytes gradually disappear these | 


imaginal oenocytes replace them; they grow quickly in size, 
and leaving the sites of formation migrate, by amoeboid 
movement apparently, into the fat-body, among whose cells 
they scatter themselves. They do not grow as large as those 
of the larva (fig. 179), seldom exceeding 2ly in diameter, 
with a nucleus of 94. The latter may be granular and may 
show several small karyosomes. Occasionally in the four-day 
pupa they may actually contain a gigantic nucleolus, contain- 
ing minute crystals; whether this is an indication of degenera- 
tion I cannot say. 

It is important to note that the oenocytes are more 
prominent in the larva than in the adult insect. This agrees 
well with the view above expressed that their function is to 
break down the storage products of the fat cells, as the 
organism needs them, feeding being ever so much more active 
during larval life. 


The Lateral Intestinal Glands. 


On either side of the intestine, just below the paired 
hepatic caeca, are to be seen, in ale mature larva, two 
organs, whose existence has not, so far as I am aware, hitherto - 
been observed in insects. The organs are in the form each 
of a long chain of very large, elongated cells about 60u in 
length, and_ presenting a weakly fibrous cytoplasm. -Within 
this delicate cytoplasm is a great heavily granular mass, oval 
in shape, and about 55u in length. It seems impossible that 
it should be anything but a greatly hypertrophied nucleus 
(fig. 169). A nucleolus may be present. 

In the larva of the first instar this organ-is_ indistinctly 
seen as a number of faintly fibrous cells just below the hepatic 


455 


eaeca; but as the larva grows they increase in size, and over- 
lapping as they grow, eventually form an elongated, well- 
defined organ on either side of the intestine. 

In the larva eight hours after defaecation a general dis- 

integration of these cells begins. In places the cytoplasm and 
nucleus may degenerate into a fine powder and be cast into 
the blood. In other places the chromatin of the great nucleus 
clumps together in numerous small balls (fig. 170), and the 
cell cytoplasm, with these degenerate chromatic globules scat- 
tered through it, floats for a time in the semi-fluid contents 
of the abdomen, and finally, sometimes by the intervention of 
leucocytes, at other times by the chemical action of the blood, 
disintegrates. At other times the apparently normal cells 
may be observed, as late as twelve, or even sixteen hours after 
defaecation, to become the prey of the leucocytes; numbers of 
these have penetrated along a channel where the hepatic 
caeca have prevented the fat-body from,encroaching too much 
upon the intestine, and here they fall upon the great hyper- 
trophied cells, and a few hours later nothing but groups of 
leucocytes, and a little débris, remains to indicate the place 
where these gland cells have once been (fig. 147, 1). 
Just what these organs are | am unable to say. Their 
structure is similar to that usually seen in gland cells; the 
absence of any duct communicating in any way with the 
intestine or any other organ, indicates that they are structures 
analogous with the various internally secreting glands so well 
known in vertebrates. | 


The Dorsal Abdominal Glands. 


These glands are to be observed in their mature condition 
only in the imago; it is not impossible that they have the 
same function here as the lateral intestinal glands have in 
the larva. 

In the early larva they are to be observed as a single 
flat band of small closely packed cells, lying upon the mid- 
dorsal region of the intestine in the hinder part of the 
abdomen (fig. 173). They show a clear cytoplasm and are 
undoubtedly in an embryonic condition. But during larval 
life they grow considerably, and separating from one another 
form a pair of long chains on either side of the heart, One 
of these is shown in fig. 213. In the defaecating larva they 
are quite large, measuring 20u in diameter. They are 
approximately spherical; their nucleus is branched, and their 
cytoplasm very vacuolated; during the remainder of larval 
life they grow a little in size, and are not unlike the 
degenerate fat cells of the late pupa in appearance (fig. 172). 


Nevertheless, they are in no way to be regarded as embryonic 
fat cells. 


456 


In the last day of larval life they proliferate; amitotically ) 
it seems, and in the larva sixteen hours after defaecation may 
extend over a considerable region of the dorsal part of the 
abdomen. Usually, however, they are confined to two chains, 
several cells in breadth, on either side of the heart. 

in this condition the cells remain during the early pupal 
period. Gradually their cytoplasm becomes more homo- 
geneous, and in the pupa shortly before emergence they may 
be observed as two irregular chains of unconnected groups of 
cells, running along the mid-dorsal portion of the abdomen. 
During pupal life the large cells as they occurred at the end 
of larval life seem to have undergone a process of incomplete 
fission, so that one now finds, not chains composed of indi- 
vidual cells, but chains of small growps of disc-shaped cells, 
arranged behind one another in little groups representing the 
cells from which they have been produced.(fig. 171). 

Within their clear, heavily eosinophilous cytoplasm lie 
numerous heavily chromatic granules which usually hide the 

_nucleus. I have observed these glands in the free-living wasp, 
“ nine days old, and there can be no doubt that they persist 
— throughout life. 

It seems impossible to regard them as anything but 
internally secreting glands. Weismann observed certain large 
cells in close connection with the heart in Diptera, and spoke 
of them as the ‘“‘cell chaplet.’”’ Lowne (1890) observed the 
same cells, and though he found them in the adult insect, 
he regarded them nevertheless as young fat cells. I do not 
know whether they are identical with the structures above 
referred to; these have, however, a remarkable resemblance 
in the immature state to fat cells. It is necessary also to point 
out that they do not constitute the pericardial septum, this 
structure being absent in the adult Vasoma. 


THE FAT-BODY. 


It is to the great development of the fat-body, together 
with the disintegration of most of the larval structures, that 
the pupa of the insect owes its semi-fluid consistency, and the 
apparent lack of organization that a superficial examination 
first reveals. 

In the newly hatched larva the fat-body is in the form of 
a number of large rounded cells, with very faintly granulated 
protoplasm and a large heavily granular nucleus, lying loose 
within the haemocoele. A single cell is usually large enough 
to occupy the greater part of the distance between the intes- 
tine and the body wall, but sometimes the cells lie ‘‘two deep.’’ 
Great gaps separate adjacent cells, and through these the blood 


457 


circulates. The fat-body is confined almost entirely to the 
thoracic and abdominal segments. 

Shortly after the feeding has commenced, the cells of the 
fat-body begin to accumulate within the cytoplasm globules 
of fat (fig. 10), and at the end of the first larval instar a 
number of these, often quite large, are present, and the cell 
has increased considerably in size, measuring now about 25u 
in diameter. In almost all the fat cells examined at this 
stage a great space was observed around the nucleus; this is 
probably an artefact. 

During larval life a great growth takes place in the size 
of these cells, till at the end they may be as large as 92 in 
diameter. This generally results in a partial crushing 
together of cells; but the increase in size of the haemocoele 
has been so great, that in places, even now, they lie loose 
within it (fig. 3). But after defaecation, when the space 
occupied by the intestine is so greatly diminished, the cells 
again separate from one another. 

In the second larval instar the accumulation of a second 
type of reserve substance becomes manifest within the fat cells 
as a heavily staining, apparently structureless, mass around 
the nucleus. But a little later this mass breaks up into 
numerous minute granules, which move partly outwards, but 
are most concentrated in the more central part of the cell. 
Other granules are formed in the more peripheral regions, 
and these are much larger, often irregular in shape, being 
sometimes even angular, and stain heavily with haematoxylin. 
The fat cells have grown greatly in size, and accumulate 
“mostly just below the cell membrane. All these storage sub- 
stances, gathered up from the surrounding blood, le sus- 
pended within the delicate, often exceedingly delicate, cyto- 
plasmic meshwork of the fat cell. 

In the larva some hours after defaecation has com- 
menced, when the imaginal discs of the integument have 
begun to grow, at the same time considerably constricting the 
body volume, the pressure exerted upon the fat cells as they 
float loosely in the blood forces numbers of these cells into 
the cavities of the outgrowing appendages—wings, legs, 
antennae—while the cavity of the head, which in the feeding 
larva was not well provided with fat cells, now becomes 
crowded with these, as the contracting abdomen presses its 
contents forwards (cf. fig. 154). 

As the abdomen contracts more and more during its 
transformation into the adult abdomen, the fat cells which 
remain within it become very tightly packed together, and 
it is only with the greatest difficulty that cell boundaries can 


458 


be detected. There is no evidence, however, that any rupture 
of the cell walls ever takes place. Between these fat cells 
lie the larval tracheoles, and the chemical disintegration 
rather than phagocytosis of these, in places which are usually 
quite inaccessible to the phagocytes, is readily understood. 

The fat cells anterior to and above the brain are in the 
form of a single layer of cells; during pupal life they are 
often to be observed showing rhythmic movements, due, un- 
doubtedly, to the contractions of the heart. The fat cells in 
the postero-ventral part of the head cavity are much more 
numerous; as in the thorax (alitrunk) they are loosely dis- 
posed and cell walls are always clearly visible. 

From the late larva till the time of death of the insect 
the fat-body undergoes a gradual degeneration and absorption, 
and, although it is quite probable that the fat-body stores | 
up reserve products as the imago feeds, yet at no time is 
there to be observed in Vasoma a formation of new fat cells; | 

| 
| 
: 
| 


the same individual fat cells which have persisted in the 
senescent imago occurred already in the first larval instar. 
It is this gradual degeneration that I shall here describe. 
In the larva at about the time of defaecation many of 
the larger grains of storage material within the fat-body begin 
to develop very minute crystals within them, and sometimes 
quite large numbers of these may be present, all within a 
single grain (fig. 92). In the small, more centrally situated, 
and eosinophilous grains these crystals are not to be seen, but 
frequently contain small chromatic granules, probably the | 
pseudonuclei of Berlese. But these crystals do not, as a | 
rule, persist long within the grain; already in larvae several 
hours after defaecation they are no longer to be seen. Even : 
the chromatic granulations of the small grains seem to dis- 
appear in the early pupa. Sometimes, however, crystals are 
visible as late as several hours after pupation. | 
The nuclei of the fat cells assume curious appearances 
towards the end of larval life. The heavily granulated 
structure and general compactness of the nucleus is lost, and 
it may become finely granular and slightly branched, while 
at other times it elongates greatly and stretches as a great 
dumb-bell-shaped band almost from one side to the other of 
the fat cell (fig. 92). At no time have I observed nuclear 


these cells as they gradually liberate their storage substances 
and then die, the processes which are to be observed micro- 
scopically are easily described. The fat globules and storage 
grains begin to decrease in number, at first slowly, then 


| 
| 
division. 
However complex may be the changes going on within 
rapidly. In the head of the three-day pupa the fat cells 


459 


have greatly diminished in size; they still contain a few 
grains and fat globules (fig. 96), but while in the adult larval 
fat cells these reserve products give the cells their charac- 
teristic appearance, the cytoplasm merely acting as a supporting 
tissue for them, in the late pupal period these conditions are 
reversed. The faintly granular cytoplasm now predominates, 
and only scattered grains and fat globules remain. The fat 
cells, however, now no longer resemble those of the first instar 
before the reserve materials accumulated. They float as 
irregular shapeless masses within the cavity of the head, and 
although a few survive the pupal period, most have degener- 
ated before then; their reserve substances have all passed 
back into the blood from which they originally came, and the 
brain and the great eyes have doubtless grown at their 
expense. 


These degenerate shapeless cells are finally, in the four- \ 


day pupa, removed si the euapeyies Sf > not before this 


degeneration, and oe. a few fat ate in rie ventral portion 
of the thorax persist, even throughout imaginal life, yet the 
greater number disappear entirely during the fourth day of 


pupal life; at the expense of their reserve substances the great 


wing-moving musculatures have developed. 

In the newly formed pupa the thoracic region contains 
numerous fat cells (fig. 154), but as the longitudinal muscles 
grow they begin to push these aside. Those cells which have 
been so unfortunate as to become entangled amongst the 
growing muscles become stretched into elongated masses, very 
well seen in the thirty-six hour pupa; the others retain their 
usual shape. But the result is always the same; the cells 
gradually give up their reserve products (fig. 98), and like 
the fat cells of the head, remain as irregular hulks, whether 
compact or branched, or greatly elongated and compressed 
between the thoracic muscles. Here in the fourth-day pupa 


they are fallen upon by the leucocytes and soon are no longer / 


seen. 

Similar degeneration may be observed in those fat cells 
which were forced into the cavities of the appendages. 
In the abdomen the degeneration during pupal life is 
much less complete; indeed, although occasional leucocytes 
may be observed lying amongst the cells of the fat-body as 
late as the fourth-day pupa, yet these seem to have no effect 
upon the fat cells. The latter remain practically at a constant 
size; the fat globules which they contain may diminish in 
number, but do not disappear. The grains, however, dis- 
appear to a large degree; the large grains are far less 


a 


460 


numerous, and the smaller eosinophilous grains are almost 
totally absent. 
Even after emerging from the pupa, the degeneration of 


abdominal fat cells continues, and it is undoubtedly at the 


expense of the fat-body that the ovaries of the female grow 
so greatly. Even after nine days, however, cells of the fat- 
body are still present in the abdomen, though considerably 
less numerous. I have not observed their disappearance, but 
it is unlikely to be different from what occurs in other places 
during pupal life. Indeed, as far as the fat-body is con- 


cerned, it is clear that the retrogressive development does — 
not cease at the time of emergence. It continues apparently — 


right throughout pupal and imaginai life. 

At times I have observed the large granular degeneration 
masses cast out by the larval integumental cells, lying 
embedded within individual fat cells (fig. 91, x); it seems, 


then, that the fat cells, in spite of their inert appearance, 


must possess a certain capacity for phagocytosis. 

The behaviour of the fat-body does not appear to be 
identical in all insects. Berlese (1901) observed multiplica- 
tion of the cells of the fat-body in the silkworm, as well as in 
certain Coleoptera. Poyarkoff (1910) observed it in Galeruca. 
In Calliphora, on the other hand, it does not appear to occur. 

Poyarkoff (1910) has described phagocytic activity of the 
fat cells of Galeruca; but it does not seem to have been 
observed elsewhere. . On the other hand, he observed also a 
phagocytosis of individual cells of the fat-body, which Pérez 
observed also in the ants (1902), and in Calliphora (1910). 
Kowalevsky (1885), on the other hand, described phagocytic 
histolysis of the fat-body in- Musca vomitaria. In living 
material he observed the leucocytes crawling over the fat 
cells, penetrating into their interior, and eventually destroy- 
ing the whole cell. It may be that this takes place under 
the influence of the egg albumen in which Kowalevsky placed 
the tissues; but no further evidence has accumulated to show 
that the phagocytosis occurs normally on an extensive scale, 
except certain observations by Lowne (1890). ‘This investigator 
described the leucocytes as entering certain fat cells, and then, 
having proliferated rapidly around the nucleus, as migrating 
outwards; the peripheral ones are much smaller than the more 


central ones, which are frequently multinucleate. The leuco- — 


cytes then leave the fat cell, which has lost, in the meantime, 
its cell membrane, and enter the blood stream. -He even 
considers the view that the leucocytes have been formed within 
the nucleus of the fat cell. Pérez could not confirm the 
observations of Lowne and of Kowalevsky, and both Weis- 
mann and Ganin, working with similar material, observed 


461 


that the fat-body disappeared only very slowly, and that many 
of the fat cells persisted even in the imago. 

In Vasoma there is a total absence of phagocytic destruc- 
tion of the food-laden fat cells. 

Several investigators have\ described a development in 
Calliphora of new fat cells for the imago. Weismann (1864) 
was the first to notice it; Berlese (1899-1901) examined the 
process more closely, and concluded that the imaginal fat 
cells were developed by the differentiation of the ‘‘spheres 
of granules.’’ This conclusion is the more remarkable when 


' it is remembered that these bodies were regarded by Berlese 


not as gorged leucocytes, but as disintegration products of 
larval cells. Henneguy (1904) adopted this view, but regarded 
the “‘spheres of granules’’ as leucocytic in nature. 

According to Supino (1900), on the other hand, the fat 


cells arise from certain mesenchyme cells, and Pérez (1910), 


in support of this view, figures a number of embryonic 
imaginal fat cells. 

_ The observation that a new development of fat cells, 
whatever the nature of the process, does occur, seems to be 
well established. In Wasonia, however, I could observe no 
indication whatever that this took place. It is perhaps useful 
to point out that the cells of the dorsal abdominal glands 
above described show a remarkable resemblance to young 
fat cells, but never develop into these. 


The Function of the Fat-body. 


Although the fat-body is a highly characteristic tissue and 
occupies so large a portion of the insect, yet its function has 
been rarely investigated, and is but little understood. It is 
beyond the scope of this paper to examine this question 
except in so far as it has a bearing on metamorphosis. 

It seems probable that the fat-body of WVasonza exhibits 


a limited phagocytic activity; Poyarkoff has seen it in 


Galeruca, and in Nasoma it appears also to be present. 

Berlese regarded the fat cells as intimately concerned 
with nutrition; food passed through the walls of the intes- 
tine, and was absorbed in an apparently solid state into the 
fat cells. Migrating inwards it came into the neighbourhood 
of the nucleus. Then it migrated outwards again, and was 
peptonised during its progress within the cell. The food was 
seen in the form of the large and small grains which are so 
prominent within the fat cells; the pseudonuclei, Berlese 
regarded, without any evidence whatever, as the enzyme, 
which brought about this hydrolysis. 

In 1889 P. Marchal observed that treatment of the fat 
cells with acids would cause the appearance of uric acid 


462 


crystals within them, and he regarded the fat-body as an 
excretory organ. 

In 1908 K. Samson observed that in the moth Hetero- 
genea the fat cells stored up vast quantities of urates during 
metamorphosis. The fact seems, then, to be fairly well estab- 
lished that the fat-body is in some way concerned with ex- 
cretion ; but whether it is a depositing place for urates, found 
elsewhere in the body, or whether the urates within it are 
the result of its own deaminising activity, these observations 
do not allow one to decide. 

In Nasoma crystals are present during late larval life, 
and a considerable portion of the pupal period, and they dis- 
appear as the urate crystals begin to accumulate within the 
intestine. Similar crystals are often seen in the nucleoli of 
degenerating larval cells, and it is possible that their presence 
within the fat cells is only secondary, their seat of origin 
being within the active tissue cells. In the larva of Nasoma, 
as already pointed out, excretory organs are absent, and unless 
nitrogen is liberated as ammonia, no removal of excretory 
products takes place. 

“Recently (1920) Pérez has shown that during meta- 
morphosis there is no evacuation of urates by the malpighian 
tubes until towards the end of pupal life. Then there is a 
sudden accumulation of urates within the intestine (just as 
occurs in Wasonza), and this coincides with a disappearance 
of the pseudonuclei from the fat-body. He regards the fat- 
body, therefore, as an “accumulating kidney.’’ 

These various investigations seem to show that the fat- 
body may remove urates from the blood during the meta- 
morphosis, and should be especially useful in such an insect 
as Nasonia, where the removal of nitrogen during larval life 
does not seem to occur. 

The fat-body has besides another great function—that of 
storing reserve materials. These are mainly in the form of 
fat globules and of the numerous grains which are so char- 
acteristic of the tissue. The latter are usually regarded, 
though without any direct chemical evidence, as protein in 
nature. 

It is this capacity of storing food materials that is so 
important in insect metabolism, and it is largely this that 
has enabled the insect metamorphosis to be evolved. 


THE GONADS. 


The Male Organs. 


The testes are present in the earliest larva as a pair of 
large pyriform structures, situated on either side of the 


463 


rectum. The narrow end of each is attached by a thin 
stalk, which is hollow, to the ventral part of the ninth 
abdominal segment, and the whole organ lies vertically to 
the longitudinal axis of the larva (fig. 185). 

The testis is covered by a membrane consisting of rather 
flattened cells—the “‘serosa,’’ or reflected abdominal ‘‘peri- 
toneum.” Lying within the sac so formed is a great mass 


of very closely packed spermatogonia, somewhat rounded cells, 


measuring about 6 in diameter. Each contains a large clear 
nucleus, the chromatin of which is concentrated into a small 


heavily staining karyosome (‘‘vesicular’’ type of nucleus) 


(fig. 186). Cell division does not appear to be going on at 
this time. 

Supporting these spermatogonia is a fine connective net- 
work, very difficult to detect; it consists essentially of a 
number of branching cells, not unlike vertebrate nerve cells 
in appearance (fig. 186), and somewhat smaller than the 
spermatogonia, the network being formed by the junction 
of adjacent cell processes. 

During larval life the spermatogonia increase in number, 
the testes in the defaecating larva being in the form of two 
rounded organs, much longer than the testis of the first larval 
instar. The spermatogonia have not increased in size; indeed, 
they are somewhat smaller than those occurring in the first 
instar, being now about 44u in diameter. The connective 
tissue network has become more prominent. 

The “‘stalk’’ of the organ, which is now definitely recog- 
nizable as a vas deferens, has increased considerably in length ; 
its wall consists of a single layer of cubical cells, covering 
which, of course, is the serosa. The lowest portion of the 
vas deferens now begins to dilate. The cells lengthen greatly, 
and change from cubical into elongated columnar cells. It 
is the rudiment of the vesicula seminalis, and is already well 


developed in the larva twelve hours after defaecation. It 


lies in close contact with the proliferating cells of the in- 
vaginated ventral part of the ninth abdominal segment, from 
which, as above described, the penis is beginning to develop. 
But its cavity does not yet possess any communication with 
the exterior. 

At this stage also (twelve hours after defaecation) the 
action of the connective tissue in the testis is clearly visible, 
resulting in the binding together of the spermatogonia in 
little groups of twenty to thirty, all clustered tightly 
together. By the time the larva pupates, these clusters of 
spermotogonia have loosened considerably; the connective 
tissue cells and network are clearly visible. Sometimes the 
connective tissue undergoes considerable hypertrophy at this 


464 


time, but this is probably to be looked upon as an 
abnormality. 

The vesiculae seminales have meanwhile been enlarging, 
and now project forwards as a pair of great thick-walled out- 
growths from the lower portions of the two vasa deferentia. 
The wasp is, then, provided with three vesiculae seminales, 
two-paired, and mesodermal in origin, formed as dilatations 
from the lower portion of the two vasa deferentia, the other 
a single forward dilatation of the cavity of the penis, as 
described more fully above (fig. 27). The cavity of the penis 


is developed about this time, and, shortly after, the lower — 


parts of the vasa deferentia open into it. 

At this time, too, the testes are beginning to elongate 
and extend forwards. The spermotogonia still measure 5y to 
6°in diameter. 

During the next twenty-four hours the male organs grow 
rapidly. The vesiculae seminales elongate somewhat and 
become ‘‘sausage-shaped.’”’ That portion of the vas deferens 
which has opened into the penis now increases in length and 

pushes the paired vesiculae upwards, so that they now come 
to lie more towards the middle of the abdomen. The testes, 
themselves, meanwhile have elongated still further, and are 
now situated dorsal to the intestine, just below the body wall. 
Their own growth, and the elongation of the vasa deferentia, 
result in their now occupying the upper regions of the fifth 
and sixth abdominal segments, having migrated through the 
seventh and eighth segments during larval and early pupal 
life. 

In the two-day pupa the openings of the vasa deferentia 
into the penis have become very wide; except for this change 
no marked alterations are visible in the male organs. The 
spermatogonia are still 54 to 6m in diameter. 

In the three-day pupa the testes fuse anteriorly above 
the intestine, and with this change, attain their mature 
proportions. 

Throughout the whole of larval, and the greater part of 
pupal life, the spermatogonia remain at a fairly constant size, 
viz., 5p to 6u. Sperm formation begins in the three-day 
pupa; I have, however, seen cases where precocious sperm 
formation took place in the pupa of thirty-six hours. The 
sperm has a rounded head about 2 in diameter; the mid- 
piece is generally quite distinct and the tail very long (about 
281). 

fe The frequent precocious development of the spermatozoa 
is especially curious; thus, while pupae three days old may 
be quite devoid of tailed spermatozoa, the pupa of fifty-six, 
and even thirty-six hours, may have testes which are abso- 
lutely crowded with sperms. 


b] 


465 


It is beyond the scope of the present paper to enter into 
any detailed account of the cytology of spermatogenesis in 
Nasomia. 

The ae Organs. 


The ovaries, like the testes, are present in the earliest 
larvae, and are not to be distinguished from these in any 


way. There is, therefore, no need to describe them here. 


Even in the larva at the time of defaecation it would be 
difficult to determine the sex of the larva, were it not for 
the presence of the rudiments of the ovipositor, and the 
absence of vesiculae seminales. The size of testis and ovary 
is fairly identical; the oogonia measure 54u to 64» in 
diameter, and, in places, are arranged in little Teer of four 
cells surrounded by a few coarse, unbranched cells, homol- 
ogous, perhaps, with the connective tissue network of the 
testis. The greater number of oogonia, however, do not 
accumulate in such masses, and the clusters are to be regarded 
as recently divided cells, which, on account of the rapid cell 
division, have not had time to separate. The oviduct is also 
a tube considerably wider than the vas deferens; proximally 
it is composed of flattened, slightly branched, cells; distally 
of cubical cells. 

In the larva at about the time of defaecation a slight 
ingrowth of cells takes place between the first and second 
pair of ovipositor appendages. In the larva, some sixteen 
hours later, this ingrowth has become more prominent and 
is beginning to undergo a certain amount of folding. It is 
the rudiment of the vagina. When first formed in the defae- 
eating larva, it consists of loosely arranged epithelial cells, 
which, however, soon merge closely together. The oviducts 
do not at this stage open into the vagina, although they 
terminate close to, and already fit tightly against the in- 
growing vaginal invagination. The ovaries have now grown 
into a pair of long spindle-shaped organs, running vertically 
and lying close beside the metamorphosing intestine; they 
reach nearly to the dorsal body wall and approach each other 
closely here, but do not, as yet, show any sign of growing 
forwards (fig. 154). 


The ovary itself consists of a great mass of oogonia, 


_ rounded or hexagonal in shape, and closely packed together. 
_ The whole mass is covered with a thin layer of minute cubical 


cells, continuous with the cells forming the oviduct; while 
covering the whole ovary is a thin serosa (fig. 187). 

A few hours after the larva has pupated the vaginal 
invagination grows backwards and begins to extend consider- 
ably in size, and the two oviducts, which for several hours 


466 


have been tightly pressed against it, now eventually com- 
municate with its cavity. At this time, also, two outgrowths 
are formed from the posterior portion of the vagina; one 
grows very rapidly and extends backwards within a few 
hours to a length of about one-third that of the abdomen. 
Already at this stage it has an extremely narrow lumen, and 
consists entirely of embryonic cells, similar to those of the 
vagina. The other outgrowth is considerably shorter, never 
exceeding half the length of its fellow. Structurally the 
two are the same at this stage; I shall speak of them here as 
the ‘‘accessory glands.”’ 

At this time, also, a pair of distinct thickenings are 


seen, one on either side of the antero-dorsal part of the vagina. — 


They will develop into the “‘lubricating glands’ of the adult 
(fig., 184). They are composed of very elongated cells, 
arranged irregularly in two ill-defined lines. 

Meanwhile the ovary has commenced to grow forwards, 
but this forward growth is accompanied by a curious parti- 
tioning of the whole ovary. The layer of small cubical cells 
covering it, and the overlying serosa begin to grow inwards 
at the tip of the ovary, in such a way as to divide the whole 
organ into four distinct compartments (fig. 192). 


As the ovaries continue to extend forwards the newly 
formed portion must likewise possess this four-chambered 
appearance. On the other hand, an extensive back-growth 
of these partitions eventually divides the whole ovary and 
even a considerable portion of the oviduct, into these four 
characteristic chambers; indeed, only the terminal portion of 
the oviduct, adjacent to the vagina, remains devoid of parti- 
tions. During the next two days the ovary grows forwards 
on either side of, and above, the intestine, and, in the 
advanced pupa eventually terminates slightly behind the 
anterior wall of the abdomen. 

In the twenty-four hour pupa, meanwhile, a new process 
of partitioning of the ovary has commenced. Ingrowths of 
the protecting membranes of the ovary divide the anterior 
tip of each of the four chambers into three secondary parts. 
The partitions do not extend deeply, but each ovary as it 
grows forwards now breaks up, as a result, into twelve 
papillae; these elongate rapidly and form twelve ovarian 
tubules, which comprise the anterior end of each ovary (see 
fig. 180). 

i The ingrowth of the external parts of the ovary becomes 
very pronounced in the oviduct of the pupa of about two 
days, being now in the form of a great connective tissue 
stroma, with four channels, each lined by a layer of flat 
cells, running along it. 


467 


The ovaries, then, so far as external appearances are 
concerned, reach their adult condition in the pupa of about 
two and a half days. Terminally each consists of twelve 
ovarian tubules, containing sexual cells, and protected by a 
thin ‘‘capsule.’”” These tubules now open into a great oviduct 
divided by a connective tissue stroma into four channels 
for the greater part of its length; but devoid of such parti- 
tions distally, near its opening into the vagina. The structure 
of the mature female is shown in fig. 180. 

The further development of the contents of these tubules, 
the oogonia, will be described below. 

It is necessary to examine now the changes undergone by 
the vagina and its accessory glands. 

In the four-hour pupa the vagina is a small sac-like 
invagination of the ventral body wall between the first and 
second ovipositor appendages; its walls consist of long 
columnar undifferentiated cells. During the next twenty-four 
hours it grows back rapidly, and extends considerably also 
in height, forming in the thirty-six hour pupa quite a spacious 
chamber on the ventral body wall, close behind the beginning 
of the ovipositor; the vagina is connected now by a distinct 
“neck’’ with the exterior. Its walls are composed of cubical 
cells; those on the upper side of the vagina, and those on the 
anterior part of the ventral surface, develop each a sharp 
forwardly pointing ‘‘tooth,’’ the inner surface of the vagina 
presenting therefore a distinctly rasp-like appearance. As 
development proceeds the cells on the upper walls elongate 
greatly, and adopt a columnar shape. A very delicate chitin 
layer is formed within the vagina, and this layer presents, of 
course, the same rasp-like appearance that occurred merely 


as a protoplasmic mould a day earlier. The function of this 


curious roughened surface is obviously to help in the laying of 
eggs. The cells themselves frequently present a clear, some- 
what vacuolated protoplasm, such as is usually seen in mucin- 
secreting gland cells. - 

On the antero-dorsal sides of the vagina a curious develog: 
ment of the epithelium has been going on, which results 
eventually in the formation of the lubricating glands, The epi- 
thelium, as already stated, consists, roughly, of two layers of 
very elongated cells ; of these cells the outer form each a gland 


_ cell; the inner, the duct of the gland cell. The outer cells 
_ Increase considerably in size, and breaking loose from the 


epithelium grow inwards a very short distance. They are 
already clearly visible in the pupa of fifty-six hours, as large 
cells with granular cytoplasm. They increase in size during 
the pupal period, and are seen in the adult insect as a pair 
of small groups of about thirty large cells on either side of the 


468 


anterior part of the ‘‘neck’’ of the vagina (figs. 180, 184). 
Meanwhile the cells of the lower layer have been differenti- 
ating. They elongate considerably, and develop, after about 
two days, a very long narrow lumen, one end of which becomes 
applied by a funnel-shaped process ‘to a gland cell, while the 


other opens into the upper part of the ovipositor. Practically 


the whole of the cell cytoplasm becomes converted into this 
duct, the nucleus itself remaining as a small sen staining 
swelling upon it. 

The function of these glands is apparently to secrete a 
lubricating liquid into the chitinous ovipositor, and aid in the 
passage of eggs down this structure, while assisting it, at the 
same time, to bore through the hard shell of the fly pupa 
during oviposition. This liquid is clearly seen during the 
act of laying as minute oily globules which ooze through the 
sheaths of the ovipositor. 

On the upper surface of the vagina two small rounded 
vesicles are seen (figs. 180, vsc.), whose walls are composed of 
long columnar cells. They appear to correspond to structures 
which in the honey bee are described as aiding in copulation, 
a kind of bursa copulatrix; what their actual function in 
Nasoma is, I am unable to say; that they have nothing to 
do with copulation seems fairly certain, since this takes place 
by the application of the penis of the male to the termination 
of the ovipositor of the female. 

The first stages in the development of the great accessory 
glands from the posterior part of the vagina have already been 
described. Two curious changes now take place in connection 
with the openings of these glands, which tend to confuse their 
true origin: firstly, the vagina grows backwards over the 
openings of the glands, so that they now arise not pos- 
teriorly from the vagina, but from its antero-ventral region ; 
secondly, shortly after the glands grow out from the vagina 
they draw a portion of the cavity of this structure after them, 
so that they open in the twenty-four hour pupa, not directly 
into the vagina, but into a separate chamber, lying beneath 
it, and opening into the ‘‘neck’’ of the vagina, shortly before 

its opening into the ovipositor. 
| The cells on the upper part of this sac elongate con- 
siderably to form a columnar epithelium; in the late stages 


of pupal life (four and a half-day pupa) their very powerful — 


staining capacity shows that they have now developed into 
gland cells. 

The two posterior accessory glands increase in length, and 
in the pupa one day old have approximately attained to their 
adult dimensions. The glands are not symmetrically placed ; 
that on the right side is much the longer of the two (fig. 180), 


i 
ee 


469 


and extends backwards to a point one-quarter the length of 
the abdomen from the posterior extermity of the insect. 
Its cells are large, and continue to develop a lumen, which 
runs right down the gland, but increases slightly in diameter. 
The cells soon lose their embryonic appearance; in the pupa 
at the end of its first day they are already wedge-shaped ; 
they have a large nucleus but present a fairly clear cytoplasm. 
In the thirty-six hour pupa, however, some of them show 
a distinct indication of developing granular cytoplasm. The 
granulations increase in number, so that in the mature pupa 
the whole cells become packed with granules; the glandular 
nature of the organ is no longer to be questioned (figs. 182, 
183). 

The gland on the left side develops into a structure 


only two-thirds the length of its fellow. Distally its lumen 


is distended into a round vesicle, and this becomes connected 
on its anterior side with a round, solid ball of cells, the spaces 
between which appear to open into the vesicle (figs. 180, 181). 


The function of these glands is doubtful. That they are 
not “‘colleterial glands’’ (glue-secreting glands) seems certain, 
for the wasp has no need to fasten her egg to an exposed sur- 
face. It is much more probable that they are lubricating 
glands, whose secretion aids that of the true lubricating glands 
in facilitating the passage of eggs down the ovipositor, and 
theentrance of the ovipositor through the hard shell of the fly 
pupa during oviposition. 

To somewhat similar glands in Calliphora, Lowne has 
applied the term ‘‘Parovaria.”’ As late as 1895 he main- 
tained, in his well-known monograph on that insect, that the 
germinal material of the egg was developed in the parovaria, 
while the yolk was produced in a pair of great ‘“‘yolk glands’”’ 
(really the ovaries), and that the large oval masses of yolk, 
as they pasesd down the uterus, first applied their microphyles 
to the opening of one of the parovaria, and received their 
germinal vesicle; then applied their micropyles to the openings 
of the spermathecae, and were fertilised. Lowne then drew 
the unfortunate comparison of the ‘‘insect vitellogen’’ with 
that of the flat worms. 


It is necessary to consider now the history of the oogonia, 
in their process of development into ova. 


Throughout larval, and the greater part of pupal life, 
the oogonia remain as small cells closely packed together, 
measuring from 5%u to 6p in diameter; each contains a large 
nucleus of the ‘‘vesicular’’ type, i.e., the chromatic material 
is contained in a minute granule, lying within a colourless 
nuclear “‘sap.’’ But towards the end of pupal life these cells 


470 


which lie in the twelve pairs of ovarian tubules begin to 
undergo a series of changes, which transform them into 
mature ova. A consideration of the nuclear changes is 
beyond the scope of this paper; I shall confine my description 
to the more obvious changes in the cells. 

The oogonia in the distal part of the tubules divide 
actively (without any centrosome, so far as I could observe) ; 
those in the proximal part of the tubes cease to divide and 
arrange themselves in little balls, which pass down the tubes 
(fig. 189) and eventually enter the four channelled oviducts. 
The grouping up of the cells into these little balls can be 
clearly observed at the point between the region of irregularly 
arranged cells and that at which the last ball has been 
formed. No difference is at this stage visible in any of the 
cells of any of these little masses (fig. 189). Very soon, how- 
ever, changes begin. The central cell of every alternate 
group begins to grow; it is the future ovum, and the sur- 
rounding cells form the follicle; the balls of cells on either 
side of these developing ova are the groups of nutritive cells. 


The follicle cells at first do not undergo any appreciable 
changes. The ovum, however, is soon characterized by a 
quickly growing nucleus. The nutritive cells soon increase in 
size ; indeed, by the time the fourth group of cells is forming, 
the nutritive cells of the first have already grown to 1llp in 
diameter. The egg meanwhile grows rapidly, but though it 
probably develops at the expense of the follicle and nutritive 
cells, these do not appear to suffer greatly; the follicle cells 
maintain a remarkable constancy in size. When the egg has 
reached a diameter (it is now slightly oval) of 12m, the follicle 
cells are still 5u to 6y in diameter; they have, however, 
become somewhat cubical instead of rounded in shape, so as 
to form a more complete covering for the ovum. 

When the egg reaches a length of 52u, the first polar 
body is formed; it is very large, measuring some 10°3p in 
diameter, and is clearly seen lying beneath the follicle cells 
(fig. 190). Even now, however, the follicle cells have not 
diminished appreciably in size; indeed, although the ovum is 
probably living partly at their expense, they may actually 
show an increase in size, reaching at times a thickness of 7p. 

The behaviour of the nuclei of the nutritive cells, how- 
ever, is quite different. The nucleus grows greatly in size 
and may reach a diameter of 5u; the chromatin is scattered 
recularly throughout it, and is no longer contained in a 
karyosome. 

Eventually, however, the follicle cells also begin to grow, 
but the growth of the nuclei never ceases. When the ovum 


re fe 


471 


measures 150, in length the nuclei of the surrounding follicle 
| cells measure 17y in diameter. Although the egg is living 
___ at the expense of the nutritive cells, these also grow greatly 
in size; it is difficult to detect their cell boundaries, but they 
show the same disproportionate growth between nucleus and 
cytoplasm, ¢.g., when the egg measures 18y in length, the 
nutritive cells measure about lly in diameter; their nuclei 
5D. 
i awe see, then, that the follicle cells and nutritive cells 
| undergo certain characteristic changes as the ovum develops; 
_ they remain of a fixed size for a time, or increase in size 
more or less rapidly, but their growth is not proportionate 
| to that of the ovum. Their nuclei, on the other hand, grow 
rapidly in size, and the rate of increase of these is much 
greater than that of the cells containing them. 
Now it has been clearly shown by Morgulis (1911) that 
the body cells of salamanders undergo during starvation a 
rapid diminution in size; also that the nuclei themselves 
become smaller, but that the rate of diminution in these soon 
becomes less than that of the cytoplasm. As a result the 
_ fatio of nucleus to cytoplasm is much greater than in normal 
| cells. Exactly how this is to be interpreted is difficult to 
_ say. It may be that the nucleus has greater powers of 
| resistance to starvation than has the cytoplasm; on the other 
hand, it seems much more correct to assume that there is an 
intimate relation between the cytoplasm and. nucleus, and that 
the condition which we find in a starved salamander cell is 
such as will enable it to exist the better under these changed 
conditions. And although this phenomenon is by no means 
universal among starving cells, still it seems to suggest that 
| a great increase in the nucleo-cytoplasmic ratio is a sign 
) that the cell is living under certain adverse conditions. 
It is in this way, possibly, that the remarkable changes 
in the nucleo-cytoplasmic ratio, undergone by the nutritive 
and follicle cells, is to be interpreted. That the nutritive cells 
nourish the ovum is universally recognized; that the follicle 
cells nourish the ovum is more difficult to prove. However, 
the fact that the latter cells undergo this same nuclear change 
is a curious piece of evidence in favour of this view. 
Considered in this light, the nutritive and follicle cells 
exhibit the interesting combined effects of nourishment and 
starvation. Their growth in size is due to their receiving 
_a large supply of nourishment; the preponderance in the size 
of the nucleus is the result of the parasitic habit of the ovum. 
The ova continue to grow rapidly, reaching at the end 
of pupal life their mature length of about 300u. They are 
_ Yeady for fertilization immediately the we emerges. 


472 


The nutritive cells, on the other hand, gradually diminish 
in size, and are left as a little clump of disappearing cells in 
close contact with the ovum. 

In the female, but not in the male, is a pair of glands 
(fig. 188) lying in close contact with the anterior extremity — 
of the ovarian tubules. They consist’ of large cells with 
granular cytoplasm, and open on to the abdomen on either 
side just behind the petiole. The glands themselves contain 
a distinct cavity. I have not observed their mode of develop- 
ment, but they seem to be formed simply as a depression in 
the ectoderm early in pupal life. 

What the function of, the glands is, is difficult to deter- 
mine. Their occurrence in the female alone indicates that 
they are sexual excitants of some kind. 


THE NERVOUS SYSTEM. 


As early as 1832 Newport, comparing the simple type 
of nervous system of the larva of Sphinx ligustri with the 
more specialized condition, with its concentration of ganglia, 
that he observed in the adult moth, showed that during 
metamorphosis a ‘‘migration’”’ of ganglia must occur: and 
examining the pupa at various stages of development, he was 
able to observe various intermediate conditions between. the 
larval and imaginal structures. 

But the first histological observations were made by 
Weismann in 1864. He showed in the muscids that a process 
of histolysis was going on within the ventral nerve cord; the 
nerve cells become dark and granular, while the whole nerve 
cord changes into a structure of very fragile consistency. The 
peripheral nerves become very pale, and losing their fibrillated 
appearance, develop fine refractile globules in their interior. 

In Corethra, on the other hand, a much-simpler process 
occurs; the central nervous system undergoes no fundamental 
changes, and only where new organs ee ah are new peri- 
pheral nerves formed. 

In 1889 Van Rees investigated the nervous system in 
Calliphora, but could not confirm Weismann’s observation 
on the fatty degeneration of the peripheral nerves. So far as 
I am aware, however, the cellular changes in the nerve cord 
and peripheral nerves have never been investigated, and even 
the work of Weismann does not contain any direct observa- 
tions on the fundamental cell changes going on here during 
metamorphosis. 

The metamorphosis of the brain has received considerable 
attention from Viallanes (1882, 1884, 1885), and much more 
recently from Bauer (1904). I shall refer to the work of 
these observers below. 


473 


The Ventral Nerve Cord and Peripheral Nerves of the 
Nasonia Larva. 


In the newly hatched larva the ventral nerve cord is 
visible through the transparent cuticle as a thick column, 
not very distinctly marked off into ganglia, and passing from 
the third segment backwards along the mid-ventral line to 
the eleventh (fig. 1). In front, the nerve cord communicates 
by a pair of circum-oesophageal connectives with the brain, 
which occupies a large part of the second segment. From the 


brain a pair of minute nerves is given off to the small rudi- 


mentary sense papillae (antennae) on the first segment. Other 
nerves doubtless leave the brain, and supply various parts of 
the head, but I have not been able to observe them definitely. 
It is only with the greatest difficulty that ganglia can be 
observed in the ventral nerve cord at this period, and the 
presence of lateral nerves is the best indication of their posi- 
tion. These nerves are quite prominent and are even clearly 
visible through the transparent cuticle (fig. 1). The posterior 
ones are the largest and supply the greater part of the hinder 
region of the larva. Lying in front of the brain, just dorsal 
to the oesophagus, is a minute rounded stomotogastric ganglion 
(fig. 117), connected by a pair of nerves that surround the 
narrow oesophagus with the circumoesophageal connectives 
near the junction of these with the first ventral ganglion. 
During the growth of the larva there is a corresponding 
increase in the size of the nervous elements, and it is not till 
a-considerable time after hatching that the various ganglia 
can be clearly observed.. Of these, twelve, not including the 
brain, are present (fig. 225, a). The last one can be seen in 
longitudinal sections to be composed, apparently, of three very 
closely fused ganglia, so that the larva possesses at least 
fifteen of these. No account is taken here of the possible 


multiganglionic nature of the brain. 


Covering the central nervous system and the peripheral 
nerves of the newly hatched larva is a very delicate membrane, 
composed of two kinds of cells: the purely larval cells, and 
the embryonic imaginal cells which will replace them during 
the metamorphosis. Both these kinds of cells are very flat- 
tened and embrace the masses of new cells closely; they con- 
stitute a part of the splanchnopleural portion of the ‘“‘peri- 
toneum.”’ 

The nerve cells are of two kinds. Lying usually on the 
outside, but sometimes also scattered partly within the nerve 
cord, are large cells, devoid of a fibre, with big nuclei, con- 
taining a large karyosome and several scattered chromatin 


i. § 


474 


granules (fig. 10). They are neuroblasts from which the adult 
nervous system will later develop. Lying more internally are 
the functional larval nerve cells, considerably smaller than 
the neuroblasts, and measuring about 44 in diameter. Hach 
has a large nucleus, surrounded by a very minute quantity 
of cytoplasmic material, all the rest of the cytoplasm being 
found in the long nerve fibres. The nerve cells are themselves 
held together by a network of neuroglia cells, usually difficult 
to distinguish from the ordinary nerve cells, but clearly visible 
in the region between adjacent ganglia. 

The nerve fibres are collected in two cylindrical nerve 
strands running along the length of the nerve cord and giving 
off branches to form the peripheral nerves in the various 
ganglia. The double nature of the nerve cord is thus clearly 
recognizable. It is usually only with the greatest difficulty 
that the individual nerve fibres can be seen, so minute and 
compressed together are they. 

The cells of the stomatogastric ganglion are similar to 
those of the ventral nerve cord. 


The Post-embryonic Development and Metamorphosis of 
the Ventral Nerve Cord. 


The cells of the nerve cord, like those of all the other 
specialized larval organs, do not proliferate, but merely grow 
in size. 

The splanchnopleural covering of the nerves and nerve 
cord may first be considered. While the embryonic imaginal 
cells do not undergo any visible changes during the larval 
period the larval cells grow greatly in size, and at the end 
of the feeding period show the usual signs of degeneration 
(figs. 221, 222), z.e., their cytoplasm becomes granulated ; the 
nuclei are long and oval, and greatly hypertrophied, 
measuring 17 in length, and contain a few scattered granules. 
The usual prominent nucleolus, so characteristic of the 
senescent cells of Vasonia, is present. 

But shortly before defaecation, the embryonic cells spring 
into activity, and diyiding | mitotically) (fig. 222) rapidly _ 
‘absorb and replace the(dying larval cells, so that several hours 
later the whole of the mesodermal covering of the nerve cord 
has been regenerated. Towards the end of pupal life some of 
the cells of this splanchnopleure develop great nucleoli, but 
beyond this no visible changes are to be detected during the 
pupal period. There is, therefore, no discontinuity inthe 
splanchopleure during its metamorphosis. Moreover, as it 
acts as a sac to enclose the nerve cells, there can be no dis- 
continuity of the nervous system as a whole during its meta- 
morphosis, whatever the changes that may be going on within 


475 


| it. These changes are very profound, and the nervous system 
undergoes as marked a metamorphosis — as does any other 
system ¢ of larval structures. 
| The increase in size of the larval nerve cells is difficult 
to estimate since most of their cytoplasm is contained in the 
long nerve fibres. In the defaecating larva, however, the 
part containing the nucleus has usually grown from a struc- 
ture which in the first instar measured about 4 in diameter 
| to one with a diameter of 43y to 5u, and sometimes slightly 
larger. The real increase in the size of the cells may be 
| judged when the growth of the great nerve strands is taken 
| into account. In young larvae these measure usually some 
| 6-10u in thickness, while in the defaecating larva they have 
grown to a thickness sometimes as much as 30u. 
| Towards the end of larval life these larval cells begin 
' to develop large nucleoli and show the typical signs of 
degeneration. 

In.the nerve cord at about the time of defaecation the 
| large neuroblasts—purely imaginal structures corresponding 
in every way with the other embryonic cells which lie dormant 
| during larval life—begin to divide by mitosis, and a consider- 
. able increase in the number of cells within the nerve cord 
| occurs (fig. 227). These cells nourish themselves, in part, | 

at any rate, at the expense of the degenerating larval cells, 
these being always recognized by their great nucleoli and 
pale cytoplasm which is in process of rapid absorption by the 
developing nerve cells. In the nerve cord the larval cells lie 
scattered among the now far more numerous imaginal nerve 
cells, and large masses of disintegrating cells are also often 
to be observed (fig. 227). In the brain this is even better 
' seen. The developing nerve cells, it seems, then simply absorb 
the dead Jarval cells, growing at their expense, and in the 


larva some twelve ‘hours after defaecation no trace of the old 
larval cells remains. 

~ As the nucleated portion of the nerve cells has thus dis- 
appeared, the long columnar nerve strand and the fibres which 
form the peripheral nerves likewise disintegrate. But the 
| appearances of degenerating nerve fibres in these two regions 
_ are quite different. 

Within the nerve cord the degeneration of the two nerve | 
strands is so intimately associated with the regeneration of 
with the highest magnifications, what is actually taking place. 
Sometimes, however, and especially within the brain, “this 

‘may be seen, the Tarval nerve strands, as the nucleated portion 
dies, begin to undergo a total disorganization, and in place of 
the strands of most delicate, almost microscopically invisible 


the nervous system that it is impossible, as a rule, to see, even | | V 


es tg 


476 


nerve fibres, we now see an irregular clumping together of the 
fibres and even partial disruption of these. Into this degen- 
erating mass now extend newly formed nerve fibres from the 
recently developed nerve cells. In the larva eight hours after 
defaecation these give off short processes, which soon become 
longer and needle-shaped (fig. 228). These then extend into 
the degenerating nerve strands as fibres of extraordinary fine- 
ness, and as the old larval fibres degenerate the newly devel- 
oped processes from the imaginal nerve cells replace them. 
These events usually run so closely together that it is not 
possible to observe either in progress; it is only when for 
some reason there is a delay in the formation of new nerve 
fibres, as often happens in the brain, that a marked globular 
degeneration of the nerve strands can be detected. 

In the peripheral nerves, however, the process is much 


more marked, and very fine instances of tissue disintegration — 


in the\absence of phagocytes can be observed. In the defae- 
cating larva, as the splanchnopleural covering of the nerve 
fibres is degenerating and is in process of rapid regeneration 


_(so that no discontinuity exists between the sheaths of the 


peripheral nerves of the larva and imago), a total degenera- 
tion of the contents of these nerve sheaths takes place. The 
constituent nerve fibres disintegrate, and the products of 
disintegration unite to form several large oval globules (fig. 
223), which, perhaps as a result of the pressure of the sheath, 
are forced along the nerve, and breaking out, evidently 
through some point of weakness, float about as small rounded 
globules in the blood stream. Here they may be in part 
absorbed by phagocytes and in part simply dissolve in the 
blood. Towards the end of the larval period the imaginal 
nerve fibres, which have been growing down and replacing the 
old larval nerve strands in the ventral nerve cord, enter the 
emptying renovated sheaths of the old larval nerves. No more 
profound tissue metamorphosis than this could be imagined, 
and yet, as far as the gross anatomical changes are con- 
cerned, no marked change occurs. It is probably these large 
elobules—degeneration products of the larval nerve fibres— 
that Weismann observed. 


Meanwhile the neuroglia network within the nervous 
system has been undergoing similar changes. This is especi- 
ally clearly visible in the larva eight hours after defaecation 
in the regions between adjacent ganglia. The larval neuroglia 
cells are observed here forming a loose network of fibres 
(fig. 229). Some of the cells are clearly in a senescent con- 
dition, presenting large nucleoli; some are growing very pale, 
evidently losing their cytoplasm, while others about them 
are growing at their expense. | 


477 


Immediately surrounding the nerve strand is a single 
layer of very small cells (fig. 227). They appear to be also 
neuroglia cells, forming a support for the fibres of the nerve 
strand which they enclose. At the time of defaecation they 
are undergoing the same changes as are taking place elsewhere 
at this period. Some of the large nucleoli are degenerating 

and being absorbed ; others are in mitosis, and are evidently 
going to replace then. pis 
| The absorption of larval cells, and the proliferation of 
h the imaginal elements within the nerve cord take place then, 
at the time of defaecation, and are complete about eight to 
_ twelve hours later. But at about the time of pupation other 
| changes which affect the gross anatomy of the ventral nerve 
cord commence. These are the changes which Newport first 
investigated, using Sphinx ligustri as his subject, and con- 
sist in a remarkable migration of ganglia, resulting in the 
‘fusion of these in groups to form the concentrated nervous 
_ system of the adult. 
_ The regenerated nerve cord is composed of twelve ganglia, 
/ connected in front by the circumoesophageal connectives with 
| the brain (figs. 225a, 231). The first ventral (suboesophageal) 
ganglion fuses with ‘the brain and will be considered in con- 
| nection with that structure. 
, A little after pupation the sixth and seventh, and also 
the ninth and tenth, ganglia fuse, so that the number of 
ventral ganglia has been reduced to ten (fig. 225c). In the 
| pupa four hours old the eighth ganglion has merged into 
the fused ninth and tenth, the number being now reduced to 
nine (fig. 225d). By the fusion of the ganglion of the pro- 
podeal (first abdominal) segment with the third thoracic 
| ganglion in the pupa eight hours of age and the absorption, 
at about this same period, of the eleventh (second last) ventral 
_ ganglion into the fused eighth, ninth, and tenth, the number 
of ganglia becomes finally reduced to seven. As the first 
ventral ganglion becomes merged into the brain it is no longer 
| recognizable as a distinct ganglion (fig. 225e), and the ventral 
nerve cord cannot therefore be said to consist of more than six 
| ganglia. In this condition we find them in the imago. The 
| first three are large and lie one in each thoracic segment ; 
connecting the last thoracic ganglion (fused fourth and fifth) 
with the first abdominal (fused sixth and seventh) is a par- 
“ticularly long nerve strand. The first abdominal ganglion is 
‘TYather small. The next two, especially the last, are much 
larger and supply the hinder and ventral region of the 
abdomen. 
i, T have not observed the cellular activities which underlie 
| these migrations of ganglia; there can, however, it seems, 


Pout 


478 


be only one process by. which this takes place, viz., by the 
amoeboid movement of the nerve cells through the fine 
neuroglia network. It is evidently in this manner that the 
cells move about within the nerve cord. 

In some ganglia the nerve cells may form layers five 
cells in thickness; these gradually diminish in number at the 
hinder and front parts of the ganglia and on the nerve strands © 
connecting adjacent ganglia may, at times, be quite absent. 

In the stomatogastric ganglion a destruction of larval 
elements, followed by a development of imaginal nerve cells, 
similar to that seen in the ventral nerve ganglia, occurs. Itis | 
unnecessary to refer further to it here. | 

It will be useful to point out that the apparent absence 
of metamorphosis in the nervous system (except for the migra- 
tion of ganglia), which is usually supposed to occur in insects, 
has never yet been demonstrated. Even Weismann’s observa- 
tions on Corethra do not wholly disprove it, the destruction 
of larval cells on a small scale being impossible to detect in 


hand dissections. - 


The most noteworthy feature of the metamorphosis of 
the ventral nerve cord is, then, the spontaneous degeneration | 
of larval cells, and their destruction not by leucocytes, but | 
by a gradual process of absorption by the growing nerve cells. 


Se 


An average sized nerve cell from the imago measures 
not more than 5p in diameter, though at times quite large 
cells, as much as 12u by 8u may be seen. The cytoplasm is 
usually much reduced, most of it having entered the nerve 
fibre process. At times a small, or rarely very large, nucleolus 
is seen. 

The splanchnopleural nerve sheath may be seen to be 
continuous, at the termination of the nerves among the 
organs, with the walls of the cells on which the nerve ends 
(fig. 226). 

The Brain. 

While it will often be possible in the following descrip- 

tion to refer to the nerve tracts within the brain, it is mani- 


festly beyond the scope of this paper to make any attempt 


to elucidate the actual nerve connections. 

The brain of the newly hatched larva (figs. 1, 230) is a | 
very prominent structure in the form of two large hemi- | 
spheres occupying the greater part of the second head seg- 
ment, and projecting forwards into the first. It measures | 
about ‘15 mm. from side to side in its broadest region, and 
is connected with the first ventral ganglion by a pair of short, 
thick, circumoesophageal connectives, which pass backwards — 


| a 


4 


479 


and downwards and enclose between them the oesophagus 
(fig. 231). 

_ The brain at this early stage is not in a very advanced 
condition, and it may be divided into two parts, an inner 
functional region and an outer region, in which active func- 
tioning does not evidently occur (fig. 230). The functional 
(truly larval) portion of the brain consists of a mass of nerve 
cells, occupying a great part of the interior of the brain. The 
_ individual nerve cells appear to be quite small, seldom more 
| than 5p in diameter ; this is due to the fact that most of their 
_eytoplasm is to be found in the long nerve fibres, whose 
' yolume it is not possible to estimate accurately. They have a 
faintly granular nucleus; the usual karyosome is very small 
' or often quite absent. 

The fibres from these nerve cells all converge to form a 
pair of great nerve tracts, one on either side, within the 
' brain, and these great nerve tracts are joined by a very 
) narrow tract from the inner portion of the antero-ventral 
‘brain region—from the antennal ganglion. Other nerve 
| fibres from this antennal ganglion unite to form a very 
minute nerve which terminates on the pair of minute sense 

papillae (true antennae) of the first segment. In this region, 
and also within the great central mass of nerve tissues, 
Synapses must occur in great numbers, but I can say nothing 
definite about them here. Some of the nerve fibres in the 
brain cross to the opposite side, others form strands which 
travel in various directions. From the brain numerous other 
fibres collect to form the two circumoesophageal nerve tracts, 
-which connect the brain with the ventral ganglia. 

Forming a distinct layer on the outside of the functional 
nerve cells are the neuroblasts, evidently non-functioning cells 
at this period of development (fig. 230). They are 8u to 9p 
in diameter, and have the ‘‘vesicular’’ type of nucleus with 
its large karyosome, so commonly found among undifferenti- 
ated cells. Though they appear to be larger than the func- 
tional nerve cells, this is in reality not so, most of the cyto- 
plasm of the latter being found in the long nerve fibres; in 
this respect, then, they form no exception to the rule that 
the functional larval cells are always much larger than the 
‘ttlon-functional imaginal cells, which will replace them during 
‘Metamorphosis. The neuroblasts form especially well-devel- 
}oped areas in certain parts of the brain: there is a pair of 
very well-defined layers, in places swollen into large masses, 
on the outer lateral regions, constituting the anlagen of part 
of the two optic ganglia (fig. 230). They extend round partly 
behind the brain as large bowl-shaped structures and give 
of forwards each a small mass of cells which projects into the 


6 —————  ——————— OC 


480 


brain amongst the larval cells, towards the great nerve tract, 
and seems to constitute the “‘bourrelet intraganglionaire” of 
Viallanes. On the internal postero-dorsal portions of the 
brain are two pairs of masses composed of rather small imaginal 
neuroblasts ; they are the anlagen of the four ocellar ganglia. 
There is another cluster of imaginal cells lying one on the 
anterior ventral part of each of the large hemispheres near 
the larval ganglia and representing the cells from which the 
antennal ganglia of the adult will later develop (fig. 230). 

During the period of active growth of the larva there is 
the usual increase in size of the purely larval elements in 
the absence of cell division. The neuroblasts do not, so far 
as I can observe, undergo any division during this period. 
But at the end of this period of activity (fourth day of larval 
life) the metamorphosis commences, at first slowly, but a day | 
later (at the time of defaecation) with apparently much 
greater rapidity. 

When the brain of the larva in the defaecation period 
is examined in sections the cells which constituted the func- 
tional part of the brain during larval life are seen to be in a 
state of advanced degeneration (figs. 77, 78,(235). The cells 
~are small and highly granular; nuclei are visible often only 
with difficulty; large nucleoli are usually present. Some- 
times the cell outlines are already becoming indistinct and | 
the whole mass is obviously in a state of active disintegration. | 
Leucocytes have not been able to penetrate to these cells, | 
and histolysis is entirely of a non-phagocytic nature. The 
great nerve tracts also show obvious signs of degeneration at 
this same time; distinct fibrillation of the tracts gradually dis- 
appears, and sometimes a faint indication of degeneration 
into fatty and other globules becomes manifest. But visible | 
degeneration in the areas, whose structures in the living state | 
is difficult enough to observe, is never so pronounced as in 
the surrounding areas where the nerve ‘‘cells’’ are dying. 

Contemporary with this extensive cellular degeneration 
a@ pronounced activity of the neuroblasts is to be observed. 
In the defaecating larva the neuroblasts have already greatly 
proliferated by mitosis, and active mitosis in various parts is” | 
still to be observed, especially in the anlagen of the optic | 
ganglia (fig. 235). And as these cells rapidly increase in | 
number they nourish themselves in part upon the dead masses — 
of nerve cells and nerve fibres, and growing gradually in bulk | 
in the larva eighteen hours after defaecation, absorb and 
replace these altogether. The cells of the two pairs of ocellar | 
ganglionic anlagen proliferate rapidly. Similar changes occur | 
in the imaginal cells (neuroblasts) of the antennal ganglion. | 
In the larva at about the time of defaecation, two kinds of ~ 


481 


dividing cells may now be clearly distinguished: there are— 
‘ (a) the large rather strongly stainifg cells which form the 
various ganglia and the ‘‘bourrelet intraganglionaire’’ 
(“Zweite Bildungsherde” of Bauer), and (6) a great mass of 
| more rounded paler cells sometimes still seen in mitosis and 
' forming the greater part of the imaginal brain where the 
' great ganglia do not occur. A small group of these is to be 
seen between the optic ganglion and the zone of degenerating 
larval cells, into which they project. It is possible that these 
constitute a mass of neuroglia cells; the proliferation of others 
which form a great ring around the bases of the two hemi- 
spheres in the neighbourhood of the ocellar and antennal 
ganglia, is resulting in a gradual constriction of the great 
central mass of degenerating nerve fibres here (cf. fig. 232, x). 

In the larva eight hours after defaecation a small mass of 
cells, often in active mitosis, appears outside the extremity 
of the degenerate nerve strand. It seems to be formed as 
an ingrowth from the optic ganglion and constitutes the 
“‘bourrelet perilaminaire’’ of Viallanes, the ‘‘(Erste) Bildungs- 
herde’’ of .Bauer (fig. 232 0.g.2). : | 

From the simple anlage of the optic ganglion three masses 
of cells therefore arise :— 


(1) Those which form what I shall call the outer optic 
ae whose fibres communicate with the compound eye. 
t is that part of the primitive anlage which remains when 
the other two parts have been formed from it, and occupies 
the position of the ‘‘optic ganglion,” as I have referred to it 
above (o.g. in figs. 80 and 236). 

(2) Those which form the middle optic ganglion, as I shall 
call it, and are represented by the ‘‘Bourrelet perilaminaire”’ 

of Viallanes. This mass first becomes visible in the larva 
i}some eight hours after defaecation (0.g.2 in figs. 80, 232, 
236). 

3) Those which will form the znner optic ganglion, and 
§)\correspond to Viallanes’ ‘‘Bourrelet intraganglionaire’’ (0.g.3 
: he figs. 80, 232, 236). 

\ 


= Ss 


ane 


eater 


ee ee 


| During larval life there is a continued growth in the 
\size of the brain, but it is in the last few hours that it begins 
to assume its adult appearance. This takes place in three 
)\ways: (a) by the gradual change in shape of the optic ganglia, 
(6) by the development of nerve fibres and synapses, (c) by 
\the gradual incorporation of the first ventral (sub-cesopha- 
geal) ganglion. 

_  lowards the end of the larval period the cells comprising 
outer optic ganglion begin to migrate outwards in the 
| meshwork of fibres formed by the perioptic membrane (see 


482 


Organs of Vision), and this gradually results in the change | 
in shape of the whole optic region, till eventually in the three- | 
day pupa the optic ganglion is mainly in the form of a short | 
stalk which connects the eye with the rest of the brain, while § 
the other two optic ganglia lie in a projection of the hemi- | 
spheres rather narrower than that in which they lay in the | 
adult larva (figs. 80, 234, 236). } 

In the early pupa, too, the antennal ganglia grow largely | 
in size (fig. 232), and form a pair of very distinct antennal | 
lobes projecting downwards, forwards, and slightly inwards | 
from the antero-ventral part of each hemisphere. The ocellar 
ganglia also project slightly on the surface of the brain. | 

In the last hours of larval life a development of the | 
nerve fibres has commenced. This is rather difficult to observe, 
for the newly formed nerve fibres grow into the degenerate | 
mass of old nerve fibres, absorbing them as they grow, and as 
the latter disappear, the others replace them, there being no / 
visible discontinuity. The only visible sign of change is | 
a gradual resumption of fibrillated appearance by the 
degenerate masses of nerve strands. The new nerve strands 
thus formed are best studied in the optic region. Between the | 
outer and middle optic ganglia such a strand, rather short | 
and thickset, and never very prominent, gradually appears, 
quite independently, in this case, of the old larval nerve | 
strand. It corresponds to the periopticon of Hickson (figs. 80, | 
236), and is formed by fibres some of which have grown | 
inwards from the outer ganglion, others outwards from the | 
middle ganglion. Synapses are doubtless formed between the | 
two. | 

The nerve cells comprising the middle and inner optic | 
ganglia likewise develop fibres, which grow, this time, | 
through the old larval nerve strand. They evidently form | 
synapses here, and the whole structure forms the second mass 
of nerve fibres, very well developed in the imago, and con- | 
stituting the ‘‘epiopticon’’ of Hickson (fig. 236). The nerve | 
cells of the inner ganglion likewise give off processes inwards 
along the old larval nerve strand, and they and similar fibres 
from more internal parts of the brain unite to form the 
‘“‘opticon’’ of Hickson—the third optic nerve strand, which 
finally brings the optic nerves into communication with the | 
rest of the brain. These changes take place in the early pupa, |} 
and so far as it is possible to observe them, are complete at 
the end of about the first day of pupal life. Many of the 
cells thus produced are true bipolar nerve cells, but many of | 
the fibres which help to form these large nerve strands come | 
from masses of cells which have not grown into the brain, | 
but have remained more at the periphery. Although I have | 


7 
: bd j 
| oF } 

d : 


| ; 483 
not been able to observe them directly, it would seem that 


the fibres ‘should be/of the T-shaped type. 
_ Meanwhile the two antennal ganglia have been increasing 
-in size. The cells in the early pupa send out processes along 


| the degenerate antennal nerve tract of the larva, and as the 


ee Sal — 


latter is gradually absorbed the fibres of the former develop 
at its expense. The fibres from the antennal nerve pass 
inwards into the antennal lobe, and within it meet and 
evidently form synapses with other fibres given off from cells 
in the more dorsal parts of the antennal lobes, these fibres in 
_ turn passing backwards and upwards as a short, thick, nerve 
| tract which enters the great irregular mass of nerve fibres in 
the middle of the brain (fig. 234). This ‘‘white matter’’ 
of the antennal ganglion is a very large and rounded mass 
_ of fibres showing shallow clefs on its surface. 
Meanwhile the great mass of paler cells described above 
has continued to grow; the cells encroach more and more 
upon the great degenerate nerve strands of the larva at the 
_ base of the two hemispheres in the region between the antennal 
» and ocellar ganglia (fig. 232x); and shortly after the first day 
, of pupal life, continuing to absorb the whole larval nerve 
* strand without proliferating, so far as I could observe, any 
' more, gradually replace this, and as nerve fibres from these 
and other cells lying more on the periphery grow into the 
dead nerve strand, this is finally absorbed and replaced by 
_ the fibres from the imaginal cells. These fibres seem to com- 
municate with others formed from the inner optic ganglion 
} and the resulting structure is the “opticon,” as Hickson has 
ealled it in Calliphora. 5 
_ By this means, then, the larval brain is gradually trans- 
formed into that of the adult. Phagocytes play no part in 
the process of absorption, but the dead cells serve directly as 
food for the growing imaginal cells. And although the pre- 
sence of mitotic figures within the brain is the only clear sign 
that development is going on at all, yet when a more careful 
| study is made it is soon seen that the brain undergoes as 
. — a metamorphosis as does any other organ of the 
ody. 

Meanwhile the first ventral ganglion has gradually become 
incorporated into the brain. In the fresh pupa, although 
_ “rejuvenation” of the ganglion has taken place, like most of 
| the other ganglia of the ventral chain, migration has not yet 
“commenced. But shortly after pupation the cells or the 
ganglion begin to migrate upwards along the circumoesopha- 
geal connectives (fig. 232). In the twenty-six hour pupa they 
have definitely become a part of the now very complex brain 
ig. 233), and during the next day they begin to consolidate 


484 


their position, and spread themselves more evenly over large 
parts of the postero-ventral part of the brain. From this 
region at least three pairs of nerves are given off to the mouth | 
parts, so that this part of the brain may be said to constitute 
a distinct lobe—the oral lobe. | 
The nerve cells comprising the brain are of the same type 
as those of the ventral nerve cord; dendrites are present, 
though usually very hard to see (fig. 224). The cells vary | 
from 34u to 7p in diameter. . 
According to the investigations of Bauer (1904) the nerve | 
cells, and even individual ganglia, are developed from single 
neuroblasts, which bud off daughter cells, which after dividing 
once become transformed into nerve cells. In WNasoma this | 
does not appear to be the case. The single-celled neuroblast | 
stage is passed through in the very early embryo, and in the 
larva of the first instar the various ganglia are already to be 
seen as distinct accumulations of embryonic cells. : 


THE VASCULAR SYSTEM. | 


(a) The Blood. } 

The blood is the great essential tissue which co-ordinates | 

the whole process of metamorphosis. It is the medium in 
which the processes of destruction and regeneration occur; into 
it the dying cells cast their products of degeneration, and upon 
its substance the growing tissues nourish themselves. 
This has been made abundantly clear in the description 

of the metamorphosis of the various organs; the actual 
chemical changes, however, which go on in the blood cannot 
be discussed here. It is sufficient to say that the globules and 
granules into which the various larval organs degenerate are 
to a large extent cast into the blood stream, where they 
dissolve. The blood, in consequence, which is usually quite 
“thin,’’ becomes during the late hours of larval life, and the | 
early hours of pupal life, very ‘‘thick,’’ and so heavily laden 
with protein materials that it often stains very -strongly in 
preparations and appears as a structureless matrix in which ~ 
the other organs lie suspended. But as the imaginal organs 
develop, these substances gradually disappear, and are no 
longer visible a day after pupation. Fl 
Frequently, however, the dead larval tissues do not have | 
time to dissolve in the blood stream; the leucocytes, instead, | 
assuming their important réle of body scavengers, fall upon | 
the dead tissues and rapidly absorb them. This - ic | 
absorption of dead tissues is very clearly seen in the removal | 
of the salivary glands, of certain tracheoles (fig. 88), of the — 
temporary pupal midgut (fig. 153), of certain muscles (fig. | 


~ 


ah 


- 


485 


105), and to a less extent in certain other tissues—processes 
which have all been described above. Nevertheless, if these 
tissues are inaccessible to leucocytes, as often happens, for 
example, through the pressure of the fat-body, then phago- 
cytosis does not occur, and the tissues undergo a gradual 
solution in the blood (fig. 91, trl.). 

Weismann (1864) was the first to observe tissue disin- 
tegration in metamorphosing insects. He regarded the tissues 
as breaking up into minute globules, ‘‘Kérnchenkugeln,’’ and 


_ to the process he gave the name histolysis. In 1884 Van Rees, 


and in the following year Kowalevsky, stimulated by Metchni- 
koff’s great discovery of the phagocytic activity of leucocytes, 


- put a special interpretation upon Weismann’s ‘‘hestolysis’’— 


they regarded it as a tissue phagocytosis, the ‘‘K6érnchen- 
kugeln” being the gorged leucocytes. 

Berlese (1901) has wholly denied the existence of phago- 
eytosis of living tissues; while Pérez (1910), working with 
Calliphora, regards the leucocytes as playing the main part 
in the destruction of larval tissues. In Wasonia there can be 
no doubt that chemical disintegration, and phagocytosis of 
dead tissues, both occur. The phagocytosis of dead tissues 
is, however, not so ingenious a device for the removal of 
débris as it at first sight appears to be; a more direct process 
would obviously be the solution of dead material in the blood. 
That this can occur in tissues which are phagocytised only 
when leucocytes have special access to them has been clearly 
demonstrated in the case of the tracheoles, and phagocytic 
histolysis is to be looked upon as the sign of a not yet fully 
eelved_metsmorphosie—ot an imperfect though ingenious 
method for_attaining a_result—which will be perfected only 
when the tissues_have ‘‘learned’’ to dissolve directly in the 
blood stream, and the leucocytes, in_their turn, to refrain 
from attacking these as they disintegrate. Berlese believes 
he has observed this kind of metamorphosis in a number of 
insects, but there is little doubt that he overlooked a quite 
extensive phagocytosis of larval tissues. It is only when other 
metamorphoses, especially those of highly specialized insects, 
are investigated, that an ‘‘ideal’’ transformation may be dis- 
covered. Phagocytic histolysis of larval tissues, indeed, seems 
to_be a very much over-estimated factor in the mechanism 
of the insect metamorphosis, and although some investigators, 
é.g., Verson, regard it only as a removal of dead larval 
tissues, others believe the leucocytes to be endowed with far 
higher powers and that they destroy the larval tissues while 
these are yet capable of actively functioning; Metchnikoff 
inclined to this view, and more recently Pérez (1910) has 
written in its favour. I shall discuss it more fully in the 


486 


second part of this paper, and shall merely remark here that 
this view is quite untenable. 


The leucocytes of the larva in its first instar are not very 
numerous; they are about 5-7 in diameter, and like most 
of the tissue cells at this stage have a fairly hyaline cytoplasm 
and a clear nucleus containing a large karyosome. They are 


still. They do not grow in size.during larval life. 

Their function during this period is apparently to engulf 
any bacteria which may have entered the circulation, and I 
have been able to observe them in young transparent larvae 
lying quite still in the blood, and absorbing minute bodies 
(micro-organisms ?) which were floating about in it. The 
are not, however, called extensively into activity till a little 
after feeding ceases. At this time the disintegration of 
larval tissues begins; and although the dead and dying larval 
cells are not bodily attacked till a day or two later, yet an 
absorption of their products of degeneration (globules and 
granules which have failed to dissolve) may occur. This 
period is marked by a great increase in the number of leuco- 
cytes, and during the next forty-eight hours their proliferation 
becomes very extensive. ; 

At first the leucocytes content themselves with absorbing 
stray granules and globules cast out by the degenerating 
cells, but later, especially at the end of the time of pupation, 
they fall upon the dead larval cells which have not yet dis- 
appeared and rapidly remove them. The process has been 
described above in the various tissues and need be only 
briefly mentioned here. As a rule, it seems, the leucocytes | 
do not enter the cells which they are attacking, as occurs so 
markedly in Calliphora, but, attaching themselves to their 
walls, send in a pseudopod, which gradually spreads out 
within the disintegrated cell, and an absorption of its sub- 
stance commences (fig. 129). It is only rarely in Vasomia that 
a leucocyte bodily enters the larval cells. At other times the 
leucocytes content themselves with nibbling off small pieces 
of tissue, which accumulate in small rod-like or rounded 
structures within their cytoplasm (fig. 195); at times, how- 
ever, a leucocyte may tear off long shreds of tissue. So large 
may these be that, to be accommodated within the leucocyte, 
they have to be bent and twisted about (figs. 197, 199). 

Division of the leucocytes appears to be only by binary 
fission. At times fully gorged leucocytes divide; but the. 
engulfed food always descends to only one of the resulting 
cells (fig. 196). I could not observe any cases of mitotic 
division. ' 


487 


The fully gorged leucocytes, which may be as much as 
ll» in diameter, begin to accumulate during the last hours 
of larval life, and the early stages of the pupal period within 
the cavities < of the various appendages. A digestion of their 
engulfed food occurs heré, and soon the food is recognizable 
only as a » number of large rounded or irregular granules within 
the cytoplasm of the leucocytes (fig. 204). Large vacuoles 
develop (fig. 194)—-structures related perhaps, in some way, 
to digestion of the granules, and these vacuoles are already 
quite commonly seen while the leucocytes are still actively 
engulfing food. Within the cavities of the appendages these 
vacuoles may increase greatly in size and considerably distend 
the leucocyte. Frequently the distension is so great that the 
leucocytes burst (figs. 205, 206), and their nuclei and degen- 
eration granules float about in the blood stream and finally 
dissolve. Usually, however, the leucocytes succeed in digest- 
ing their meal and gradually diminish in size again. The 


granules slowly disappear, and only small vacuoles remain 


(figs. 208, 209). In this condition they persist throughout the 
life of the insect. They are about 7y in diameter, and have a 
large nucleus, with a faintly granular chromatic content, and 
one small | karyosome. 

(0) The Heart .”” 


The fate of the heart during the insect metamorphosis 
has, so far as I am aware, never been carefully investigated, 
and the most contradictory views are held as to the events 
that occur within it during this period. 

According to Newport, the dorsal vessel of Sphinx 
hgustri pulsates throughout the whole of the pupal period, 


and evidently undergoes no changes during this time. In ‘\ 


Eristalis, on the other hand, Kiinckel d’Herculais observed a 


day of pupal life. 


According to Kowalevsky (1887) the dorsal vessel in 
Calliphora pulsates regularly till the third day of pupal life; 
thereafter it beats more irregularly, but does not seem to 
undergo any metamorphosis. Lowne (1890-1895) partly in- 
clines to this view; he observed a change in the form of the 
heart, but attributed this to a possible replacement of its 
old muscle cells by new ones. 

Weismann (1864), with no modern technique available 
to him, came to an entirely different conclusion. He observed, 
in hand dissections (!), that the heart became more fragile and 
was evidentiy at this time in a state of ‘‘histolysis.’’ ‘‘As 
an organ it is not broken up, but is redeveloped by a process 
similar to that which has been observed in the intestines and 
malpighian vessels.’ 


i / 
} a 
| on Os 


cessation of the heart beat for one or two days after the eighth | 
a 


488 


If the transformation of the heart of Vasonia is any indi- 
cation of what happens in the blow-fly, then there can be no 
doubt that Weismann’s view was the more correct. In the 
heart of NVasonia a metamorphosis occurs quite as profound as 
that observed in any of the other organs. 


The Structure of the Larval Heart. 


The heart of the larva (fig. 211) is a long tube, running 
right along the mid-dorsal region of the body and gradually 
bending downwards near the middle of the body, terminating 
in front, close behind the brain. It measures about 1°6 mm. 
in length. It is widest behind, where it measures ‘(08 mm. in 
diameter and gradually tapers in front into a long capillary 
tube—the “‘aorta.’’ 

The heart lies within the pericardium, a tube composed 
of a fine delicate membrane (fig. 213), which opens just behind 
the posterior part of the heart by a large funnel-shaped open- 
ing (figs. 211, 212), and tapers gradually anteriorly, eventu- 
ally fusing in the anterior part of the body, with the heart. 
The pericardial walls are composed of a single layer of greatly 
flattened cells (fig. 213), but are quite devoid of muscles. 


While the pericardium has a wide funnel-shaped opening 
into the body cavity behind, the heart itself is closed, and has 
no communication with the pericardial cavity except by a 
series of six pairs of minute openings, the ostia. These ostia 
are usually very difficult to observe, the only prominent ones 
being a single pair at the posterior end of the heart (fig. 212). 
The cardiac walls are here deflected inwards to form a valve, 
which allows blood to flow only from the ‘pericardial cavity 
into the heart. 


Neither the walls of the pericardium nor of the heart 
are themselves contractile; pulsation of the heart is produced 
by the contracting of certain very delicate muscles inserted in 
_the walls of the heart in irregular pairs at intervals along its 
length; the other ends of these minute muscles are inserted 
directly on to the dorsal integument. The pericardial walls 
are drawn out into long conical processes, within which these 
muscles lie (fig. 211). So far as I could observe, the peri- 
cardium is itself not contractile, and alary muscles appear to 
be quite absent. The pericardium, then, appears to be different 
from what is supposed usually to occur in insects. 


The cells of the heart undergo the same changes during 
larval life as do those of the other specialized larval organs; 
there is a great increase in cell size, with a total absence of 
cell division. In the adult larva they have attained a great 
size (fig. 216); they are very flat and measure as much as 


2 


489 


34u in length. The nucleus is rounded or seat measuring 
5u by 12p, and has a small nucleolus. 


The Metamorphosis of the Heart. 


It is at about the time of defaecation that the heart 
begins its transformation, and the metamorphosis is very pro- 
found. In the period just prior to defaecation the pericardial 
and ‘heart cells begin to undergo a granular degeneration. The 
nuclei show the usual great nucleoli, and the cells the usual 
hypertrophy (fig. 214). At this time it becomes possible to 
distinguish clearly the larval from the imaginal elements 
within the heart tissues (fig. 213). The imaginal heart, how- 
ever, is regenerated not only from.scattered embryonic cells 
within lin its walls, but also from a mass of embryonic cells lying 
just below_ the heart, and dorsal to the rear of the midgut. 
I have not been able to locate this structure in the early 
larva, but at the time of defaecation a column of these 
embryonic cells may be observed extending upwards and along 
the ventral side of the pericardium. Proliferation is very 
rapid and the cells advance quickly, absorbing the larval 
elements as they grow (figs. 214, 215, 216). The heart itself 
is rejuvenated mainly or entirely from the imaginal cells 
within its walls; only the pericardium seems to arise from the 

“‘sub- pericardial imaginal disc.’”’ In the larva eight hours 
after defaecation the heart tube has been completely rejuven- 
ated, and below it, and in close contact with it, lies a long 
band of cells, the renovated | “pericardium” (fig. 217). The 
nuclei of the (true) heart cells are large and bulge into the 
lumen of the tube (figs. 217, 219). At this period the 
imaginal pericardial cells are in process of rapid division. 
They quickly grow upwards (fig. 218) and soon form another 
tube on the outside of, and in close contact with, the 
renovated heart. The pericardium, therefore, no longer 
forms a loose sac on the outside of the heart, but actually 
becomes a part of it (fig. 219). In the region of the stomach 
the ‘“compound”’ heart remains as a rather wide sac-like tube. 
Ostia become developed, but I have not observed their dis- 
position accurately. In this region, also, the ‘‘pericardial’’ 
cells, which have formed a membrane in close contact with 
the 1e true heart tube, transform themselves into striated muscle 
cells (fig. 220). “This part of the heart alone is contractile. , 


All the more anterior part (7.e., in the thoracic and anterior ~“ 


abdominal regions) is to be looked upon as an “‘aorta.’’ It is 


composed, of course, of the ordinary heart tube, and the 


surrounding closely fitting renovated pericardium (fig. 219). 
The dorsal vessel of the adult wasp consists, then, of a 
short contractile chamber lying in the posterodorsal region of 


. 
: 


490 


the abdomen; it alone is contractile, and is not surrounded 

_ by pericardium. Its walls consist of two layers, an inner 
exceedingly fine and delicate ‘‘endothelium,’’ the transformed 
larval heart; and an outer layer of broad striated muscle 
fibres, circularly disposed, corresponding and homologous with 
the pericardium. The aorta, which is continuous with it in 
front, gradually bends down, and lies just dorsal to the 
oesophagus (cf. fig. 154). It is, of course, composed of the | 
same two layers (fig. 219). 

During larval life a few large rounded cells le in clos 
connection with the heart. They resemble the cells of the 
dorsal abdominal glands in appearance. So far as I could 
observe, they disappear late in pupal life. 

As the thoracic intestine, some six hours after pupation, 
begins to assume its almost insignificant proportions, the 
anterior part of the heart (aorta) sags downwards, and comes 
to lie just dorsal to the oesophagus, 7.¢., a little below the 
mid-region of the thorax. 

I have not been able to observe ne heart-beat during 
metamorphosis, the heart being too obscured by the surround- 
ing fat-body. In the head of the pupa, however, movements 
of the fat-body regularly occur, and this is due, doubtless, 
to the beating of the heart. 


A ppendix. 
THE DEGENERATION PROCESSES OF THE 
LARVAL CELLS OF VASONTIA. 


The physiological interpretation which we place on the 
insect transformation depends in the main on our knowledge 
of the condition of the larval cells at the time of meta- 
morphosis. Do the cells die, and is it only dead material that 
the leucocytes absorb; or are they attacked by the leucocytes 
while still alive? It is the former opinion, to a certain extent, 
that Berlese holds: ‘‘Phagocytosis never occurs, and amoebo- 
cytes only become active when the muscle has disintegrated 
through internal causes.’’ His amoebocytes were, mainly, ab 
any rate, embryonic cells—‘‘Myocytes,” ‘‘Splanchnocytes,’’ 
etc.—but there can be little doubt that his ‘‘sarcolytes’’ were 
really gorged leucocytes, and that phagocytosis does occur is 
certain. But is it phagocytosis of dead or living larval cells? 

Pérez has written in favour of the latter view. ‘‘I think 
I have proved satisfactorily that there is no spontaneous 
fragmentation of this organ into sarcolytes, as Berlese 
thought.’’ He observed the phagocytes entering muscles 


“J 


491 


always of apparently normal structure. But at other times 
he was clearly dealing with degenerating cells—cells with 
globulated and highly vacuolated protoplasm, though the 
occurrence of large vacuoles in the living salivary glands led 
him to doubt this interpretation. It should be pointed out, 
however, that even if a cell still has its normal structure, that _ 
is no proof | that it is snot dead. It may take many hours for 
the Se achiral “symptoms of death to become visible. It is, 
perhaps, just necessary to add that in whatever condition 
the tissues may have been when they were fixed, there is no 
doubt that when they were examined they were dead. It is 
also worth drawing attention to the fact that Lowne has 
expressly stated that the larval muscles become functionless 
some time before phagocytosis commences. The muscles of 
Calliphora appear to be very resistant to autolysis; torn-off 
fragments engulfed by phagocytes still retain their normal 
structure, yet these are, it is to be presumed, quite ‘‘dead.”’ 
When, however, we examine the cells of the adult 
larva of Wasomia no doubts can be left in our minds 
that death, accompanied this time by active disin- 
tegration, has occurred, and the most varied types of 
disintegration are to be seen. _AJI] the cells show as 


a common feature _a great hypertrophy. They have often 


grown to many times the size of the cells of the newly 
hatched larva; even the nerve cells have grown greatly, though 
here the increase in size cannot be estimated, as the volume 
of the nerve fibres is indeterminable. Most of the degener- 
ating cells present, also, a great nucleolus within the nucleus. 
Sometimes this may be relatively gigantic and may lodge 
excretory(?) crystals. This in itself lends support to the view 
that the nucleolus is a structure within the nucleus concerned 
with excretion—perhaps itself an excretory product, perhaps 
an excretory “‘organ.’’ 

The disintegration of the various cells occurs, as we have 
seen, In many ways. Sometimes before this -has had time to 
proceed very far, phagocytes or embryonic imaginal cells may 


overwhelm them (many muscles). At other times the adjacent 


imaginal cells absorb the degeneration products of the dead 
larval cells directly, either by secreting an enzyme which dis- 
solves them, or by waiting for them to disintegrate spon- 


taneously (microscopic examination cannot decide between 


these two). This is seen in the nerve cells. 

But at other times the larval organs undergo most marked 
visible disintegrations.. The cytoplasm becomes disorganized 
and may break up into globules or granules, or into a very fine 
débris, and be cast, as one large globule, or numerous minute 
particles from the cell, the wall of which itself later dissolves 


492 


in the blood stream or is absorbed by phagocytes. One of the 
commonest sights in the pupa is the hulks of these old cells, 
usually quite devoid of any trace of cytoplasm or nucleus, 
floating about in the blood stream, waiting to be engulfed 
by phagocytes. Sometimes the muscles may disintegrate 
spontaneously and may break up gradually into small globules 
which break through the sarcolemma and dissolve in the blood 
or become phagocytised; but perhaps the most extraordinary 
case of spontaneous disintegration is seen when the whole of 
the minute rod-like sarcous elements, which comprise the 
striations, are cast as a shower of minute particles into the 
blood, where they gradually dissolve (fig. 104). 

At no time have I ever observed the phagocytosis of 
tissues which could be regarded as living, and even in those 
cases where embryonic histoblasts overwhelm the larval organs 
and develop at their expense, visible degeneration has always 
previously occurred. 


Part II. On the Physiology and Interpretation of the 
Insect Metamorphosis. 


The constant occurrence of so many different characters, 
often of the most trivial kind, amongst even the most widely 
separated orders of insects, is in itself sufficient evidence 
to show that the ‘‘insect type’’ must have been evolved before 
the many varied kinds of development which we see at. the 
present day amongst insects, existed. It is inconceivable that 
such organs as compound eyes, wing nervures, insectan mouth 
appendages, legs of a constant character, to mention but a few 
of them, should have been produced again and again from 
independent sources. 

Nor is it difficult to decide whether these primitive insects 
showed the direct or the more complicated type of develop- . 
ment. The oldest insects yet discovered—the Palaeodictyop- 
tera and Protorthoptera of Carboniferous times, were clearly 
related to insects which at the present day show no meta- 
morphosis. The discovery in the Permian rocks of Russia of 
an ephemerid type of larva shows that already at this early 
period the indirect type of development, though as yet not 
of a very profound nature, had begun to evolve. There can be 
little doubt, however, that when the insects which must have 
existed in the earlier Palaeozoic ages become known to us, 
none but the most generalized of types will be found to have 


493 


existed. It may, of course, be said that insects of a kind 
which now undergo no metamorphosis may have done so at 
an earlier period; there is, however, no need to make this 
assumption, for, as I shall show later, the evolution of meta- 
morphosis has been a necessary consequence of the specializa- 
tion which these early generalized orders have since undergone. 


In order to trace the stages through which metamorphosis 
has been evolved it will be necessary to describe, very briefly, 
the’ main features of the postembryonic developments of a 
number of insects, in so far as we know them. As many 
of the accounts are not very reliable, and as no metamorphoses 
have been investigated from this point of view, the comparison 
is less complete than it ouglit to be. The insects are arranged 
in what I regard as ascending degrees of metamorphosis. The 
reason for this will be clear later. It is very interesting, at 
the same time, to observe that the order is also approximately 
that of increase of specialization. 

(1) The Aptera.—These emerge from the egg in practic- 
ally the adult condition. In Machilis the eye is believed to 
continue to develop. Sexual organs undoubtedly ripen during 
post-embryonic life. 

(2) Orthoptera and Hemiptera.—These usually emerge in 
a condition which, though like that of the adult, is somewhat 
more generalized, e.g., the thoracic segments have not yet 
markedly differentiated. During post-embryonic life there is 
a gradual growth of the wings; the insect moults several 
times, but only at the last moult do the wings appear free on 
the surface. Sexual organs undergo a parallel development. 
In the wingless forms post-embryonic life is of the .apteran 
type. 

(3) Odonata.—Partially developed wings clearly visible 
through the integument, even in early instars. Legs always 
very well developed, resembling those of adult. ‘‘The internal 
metamorphosis begins considerably before the hatching of the 
insect; the larva refuses to feed and is restless. Hypoderm 
cells proliferate, causing the larva to appear tense and swollen. 
The wing muscles grow greatly and increase the size of the 
thorax. New elements form rapidly in the eye.’’—Tillyard 
(1917). The larva then leaves the water, and the moult dis- 
closes the adult insect. Nothing is known of the cellular 
changes underlying the process, but they are evidently of the 
highest interest. : 

(4) Coleoptera (type: Galeruca, examined by Poyarkoff). 
—Larvae emerge with typical insect head and mouth 
appendages. Legs present, but much more reduced than in 
Odonata. Wings never clearly seen in larva. Division of 


494 


purely larval cells occurs during the feeding period, and there 
is a great growth in size. At metamorphosis some of the more 
specialized tissues (muscles) are phagocytised (evidently after 
dying); the cells of the less specialized tissues (hypoderm, 
oesophagus, rectum) cast out parts of their substance, and 
having evidently rejuvenated, remain as the adult tissues. In 
the case of the other organs, those of the imago are formed 
from areas of cells which till now have lain dormant—the 
imaginal discs. 

(5) Lepidoptera.—Larvae emerge with typical insect head 
and mouth appendages. Legs present though very reduced. 
The larval cells divide just before the various moults during 
_ the larval period. The larval tissues seem to disappear by 

phagocytosis and redevelop from imaginal discs. Very little 
is known, however, about the Lepidopteran metamorphosis. 

(6) Muscidae (Weismann, Kowalevsky, Van Rees, Pérez, 
and others).—The larva emerges as a legless maggot, with 
reduced spiracles, and with poorly developed mouth append- 
ages. Except for a tracheal proliferation cell division does 
not occur during larval life. At the end of the feeding period 
the larval cells die and are phagocytised; the imago develops 
from imaginal discs of scattered imaginal cells. 

(7) Chaleid Wasps (type: Nasoma).—The larva hatches 
in a very primitive condition; the head is still in a biseg- 
mented state; the only mouth appendages present are rudi- 
mentary mandibles; malpighian tubes have not yet developed ; 
the proctodaeal invagination has not yet even opened into 
the archenteron. There is no cell division during larval life, 
except possibly in the tracheal system (most of the apparent 
proliferation here is, however, due to growth in cell size). 
After three days all the larval tissues—even the almost in- 
significant peritoneal membrane—having grown in cell size, 
die. Some are removed by phagocytes, others dissolve in the 
blood. The imago develops from imaginal discs or scattered 
imaginal cells. 


It seems to follow from the above account that a meta-_ 
morphosis—a period of more or less violent transformation in 
contrast with ‘‘normal’’ progressive development—occurs in 
its simplest form in the dragon-flies. No definite pupa is 
formed here, and the insect, though restless and refusing to 
feed, is never helpless. But in all the others the internal 
changes are of a much more violent nature, amounting at 
times to an absolute death—in the fullest sense of the word— 
of the larva. If imaginal tissues were not present the 
‘corpse’ would be left to decompose; the imaginal’ cells, 


owe 


495 


however, take the opportunity, and nourishing themselves 
upon the highly nutrient material of the dead larva, grow by an 
orderly process of development into a totally different organism 
—the imago. Even the instincts of the metamorphosing 
larvae are those of dying animals—they avoid the light; they 
roll themselves up in leaves; they crawl into secluded spots, 
or, significantly, even bury themselves in the earth! And yet 
by a wonderful process of development, unique of its kind 
among living things, the dead larva becomes the prey of 
minute scattered cells which have lain helpless among the 
larval cells while these flourished ; having awaited their oppor- 
tunity, these now spring into activity, and nourishing them- 
selves upon the. bodies of the dead larval cells, form another 
organism. From the grave of the dead larva arises the perfect 
insect. 

While then in the simpler metamorphoses there is a rapid 
transformation of tissue, in the more profound type a serious 
disruption of tissues occurs, the embryo developing within 
the old larval sheath is in a ‘helpless condition, and we speak 
of it as the pupa. 

The metamorphosis of an insect consists, nhl ble of two 
processes—a process of disruption and a process of orderly 
embryonic development. A ‘‘complete’’ explanation of meta- 
morphosis will, therefore, have to account for the disruption 
and likewise for the orderly development which ensues. The 
mechanism of the development is identical with that of any 
other embryonic development; it is the unexplained ‘‘Ent- 
wicklungsmechanik der Organismen,’’ and I can say nothing 
of it here. For-the process of disruption—of actual trans- 
formation—however, a simple explanation may, I think, be 
given. As it is highly probable, moreover, that the factor 
which brings about disruption of larval tissues in the more 
complex metamorphoses is identical with that which forces 
the cells in a simpler type to rejuvenate, it should be possible 
to obtain a general principle underlying metamorphosis. And 
lastly, since the rejuvenation of individual cells is itself evi- 
dently a process of rapid differentiation in cells in which the 
process has for some reason become temporarily suspended, 
it would seem that the same principle should be responsible for 
the cell differentiation seen in ‘‘normal’’ embryonic develop- 
ments. The discovery of the mechanism of metamorphosis 
ought, therefore, to lead to a more general theory of cell 
differentiation. 

As the principle is most clearly revealed in the more pro- 
found metamorphoses such as that of Vasonia, or the Muscids, 
these will be considered first. In the section on cell degenera- 
tion in the larval tissues of Vasoma I have pointed out that 


' 


( 
\ 
} 


496 


phagocytic histolysis consists always of a removal of dead 
tissues by leucocytes or their absorption by embryonic cells; 
if these cells do not intervene, the dead tissues will dissolve 
“of their own account in the blood. All attempts to explain 
metamorphosis, therefore, which have concerned themselves 


/ with the phagocytosis of /iving tissues, have proved unsuc- 
' cessful. Metchnikoff, for example, in seeking to explain the 
' immunity of the imaginal cells at a time when the larval cells, 


which had but a day before been in the height of their 
activity, were becoming overwhelmed by leucocytes, concluded 
that the imaginal cells must emit substances which held the 
leucocytes at bay, and that the larval cells at metamorphosis 
no longer did this. He was led to this conclusion by his 
belief in the existence of anti-leucocytic substances in virulent 
anthrax bacteria which were not phagocytised. . 

Of the death of the cells before phagocytosis there can, 
however, be no doubt. While this view has more than once 
been favourably accepted, the cause of the extensive cell-death 
has not, so far as I am aware, been satisfactorily explained, 
and this, after all, is the real mechanical principle that under- 
lies metamorphosis. It is usually believed that the extensive 
cell-death has been produced by the ‘‘wearing out” of a great 
number of cells all at one time. Such wearing out of cells is 
believed to occur also in other organisms, but here it is 
gradual, and as some cells die others replace them. The latter 
must then be the imaginal cells of the metabolic insect, and 
metamorphosis has been evolved by the dying cells all ‘‘learn- 
ing’’ to undergo senescence at one moment; this has evidently 
been evolved in response to the necessity for a metamorphosis 
in animals whose young and adult stages have different feed- 
ing habits (Lubbock). 

Now this explanation is clearly not very satisfactory; if 
the extensive cell-death is merely a concentration, at one 
moment, of the deaths of numerous cells which would normally 
take place gradually throughout the life of the insect, how 
are we to explain rejuvenation of cells in metamorphoses of a 
simple type? Metamorphosis by cellular rejuvenation would 
rather appear to be a stage intermediate between a simple 
development and a total disruption followed by redevelopment 
from imaginal cells, as seen in the more profound meta- 
morphoses. Moreover, the essential thing to show would be 
how this concentration of death points has been produced, 
and this is manifestly beyond the scope of modern cytology. 

The extensive tissue disruption, it seems to me, is to be 
explained on a much simpler principle. The larva consists, 
in the highly specialized forms, of very specialized cells. Such 
cells never proliferate, and the growth of the larva is due 


497 


entirely to a growth in the size of the larval cells. Now as 
these cells grow larger and larger there is formed an increasing 
disproportion between the volume of the cell contents (which 
face membrane through which the cell contents are being fed 
(and which increases only as the square of the radius). 
Eventually, therefore, a time must come at which the cell 
contents cannot any longer receive sufficient nourishment 
through the.cell membrane. Death by starvation must be the_ 
result—indeed, the more rapidly y the larva feeds, the sooner 
it will starve. This great increase in the size of cells must 
also have another effect. The delicate chemical reactions 
occurring within the cells are of such a nature that they are 
very efficient in minute cells; but it is quite conceivable that 
as the distances through which diffusions and other molecular 
movements, taking place within cells, become increasingly 
larger, a critical volume will be reached at which the delicate 
balance between the reaction is upset and cellular death is 
the result. As the cells all grow approximately equally rapidly, 
a simultaneous death of cells throughout the larva will occur ; 
the leucocytes then fall upon the débris, and the result is 
phagocytic histolysis. In insects which do not undergo meta- 
morphosis, on the other hand, growth is produced largely by 
cell proliferation. The cells do not reach their critical state 
and extensive cell-death does not occur. 

While I am convinced that it is nothing but this great 
cell growth to which the whole wonderful transformation is 
to be attributed, yet I am aware that a number of objections 
may be made against it. If, as it seems, the individual cells 
have certain critical values beyond which life is no longer 
possible, how is it that underfed larvae, in which the cells 
have not reached this critical volume, may nevertheless meta- 
morphose? It is to be noted, however, that in underfeeding © 
(partially starving) the active larvae we are actively applying 
the very factor which must inevitably appear as the cells 
hypertrophy, vz., starvation. This objection may, indeed, be 
turned into a strong support for the view which I have above 
expressed. 

A more serious objection is that in cold weather the 
larvae may live for months after the cells have attained the 
critical volume. However, it has been pointed out by Chun . 
that the abundance of life in arctic and antarctic waters is 
due to the greater length of life of the organisms living there, 
due to a slackening of the metabolic processes with the lower- 
ing of the temperature. Probably the lowered temperature 
acts similarly on these mature larvae, temporarily repressing 
those chemical reactions within the cell which result in cellular 
disorganisation. 


498 


The extensive tissue disruption which occurs at meta- 
morphosis is due, then, to the hypertrophied state of the 
cell, this in turn being the result of a failure of the larval 
cells to divide. A more complete explanation ought, then, 
to account also for this absence of cell division with the devel- 
opment of specialization by the larval cells. Now I have 
pointed out, in the foregoing account of the metamorphosis of 
Nasoma, that the larval nucleus does not seem to be able to 
increase its chromatic contents. A volumetric: increase— 
sometimes quite considerable—of the nucleus is frequently 
seen, but no increase in the quantity of chromatin is ever to 
be detected. There may be an increase in the number of 
chromatin granules, but these are always formed by a break- 
ing up of the karyosome of the nucleus of the young cell 
(fig. 102). Now it has been pointed out by Professor Brails- 
ford Robertson (1909) that cholin formed as a by-product 
during the synthesis of nuclein from lecithin, accumulating 
mainly at the median plane of the cell, would bring about a 
diminution of surface tension here, and division would result. 
In the growing larval cells such nuclein synthesis is apparently 
absent, and consequently no cell division is to be expected. 
Specialization may perhaps consist then, in part, simply in 
the loss of the nuclein synthesizing enzymes. 

The investigations of Poyarkoff have shown that in 
Galeruca many of the larval cells transform themselves into 
imaginal cells by undergoing a process of rejuvenation; por- 
tions of the old cells are cast out and the interactions of the 
remaining substances transform the cell into that of the adult. 
The cytological interpretation of these results is very difficult. 
But is it not possible that those cell substances which have 
(phylogenetically) recently been acquired—substances the 
acquisition of which has enabled the cells gradually to adapt 
‘themselves to the new conditions, as the feeding habits of. the 
evolving larva gradually diverged from those of the adult— 
may become starved out from the cells as they gradually 
approach the critical volume? These substances are, in a 
sense, ‘‘foreign’’ to the cell; while the cell lived under its 
new environment (the /arval environment) they thrived; but 
as the cells gradually weakened with increasing cell volume, 
would it not be these very substances which would perish first ? 
And is it not possible that the substances which Poyarkoff 
observed emerging from the rejuvenating cells were nothing 
but the substances to which the larval cells owe their new 
properties? These considerations will become clearer when 
we have examined the phylogeny of the insect metamorphosis. 
It is sufficient to point out here that even in the ‘‘cell 
rejuvenation’’ type of metamorphosis the same stimulus—the 


499 


attainment of a critical cell volume—may bring about the 
sudden transformation. In these forms divergence of the cells 
from the imaginal condition has not proceded so far as to 
prevent the cells returning to it; but in the more profound 
metamorphoses the greater specialization of the larval cells 
has resulted in a far more marked departure from the imaginal 
type; the cells are unable to recover when they reach the 
critical volume, and death is the result. 

Cell rejuvenation, then, is to be looked upon as a sudden 
differentiation in a cell in which this has been, for some 
reason, delayed ; it differs from differentiation as more usually 
seen, in that here the process is gradual. No satisfactory 
explanation has yet been offered for the extraordinary 
phenomenon of cell differentiation—the gradual transforming 
of a cell from a non-differentiated truly embryonic state into 
one whose structure is correlated with its function. But the 
“‘abnormal’’ differentiations to be observed in metamorphosing 
insects seem to throw considerable light on the process. It 
is well known that non-differentiated cells may quite success- 
fully perform work which is usually carried out by differenti- 
ated cells. For instance, the heart of a chick embryo beats 
long before cardiac muscle becomes .differentiated. I have 
similarly observed undifferentiated muscle cells of NWasonia 
functioning successfully. Now, it is well known that cell 
growth and cell differentiation are parallel events in embryonic 
processes. ‘Tissues consisting of cells with definite hereditary 
characteristics are laid down; the constituent cells grow and 
the tissues, or better, their component cells, differentiate. 
Is it not possible that in the struggle for existence that must 
ensue as the cells grow in size, all but the non-essential sub- 
stances—all those substances to which the generalized con- 
dition of the embryonic cell is due—gradually disappear ? 
Only those substances which are essential persist. The 
“explanation”’ is very incomplete, and there are many diffi- 
culties in the way of its acceptance. But it seems to me that 
it contains an element of truth. 


It is necessary to consider next the course of evolution 
of the insect metamorphosis, and to consider the factors 
which have necessitated this evolution. Lubbock has pointed 
out that a metamorphosis is a necessity in organisms whose 
adult and larval mouth parts are structurally unlike. This 
does not, however, explain the reason for the metamorphosis 
of the more insignificant structures of the insect’s body. Nor 
does it help us to understand why the insect larva which shows 
this metamorphosis should ever have been evolved. 

The phylogenetic significance of the larva has, I believe, 
frequently been explained correctly. It is a stage which has 


500 


been gradually inserted into the direct development; feeding 
gradually became confined to the earlier part of the life 
period, further development to the latter. In other words, 
processes which ran side by side, have gradually become 
separated. This conception is undoubtedly correct; but no 
explanation has, so far as I am aware, ever been offered as to 
why such a complex’ process should ever have replaced the 
simpler one, nor have the various types of metamorphosis 
ever been considered as throwing light on the structural 
changes through which the insect passes as they developed 
metamorphoses. It is these questions that I wish to discuss 
here. Ae 

When we consider the insects as a group and seek to 
account for their extraordinary success in nature, two char- 
acters in their structure present themselves to us—the wings, 
which have enabled them to conquer a new environment; 
and the hard chitinous body wall, which makes them so secure 
against attack. For the less specialized insects—cockroaches, 
silverfishes, grasshoppers, etc.—which live in truly hidden 
localities (Cryptozoa), these structures, though important, are 
not constantly essential. A grasshopper may pass much of its 
life without wings and ‘may even temporarily cast its cuticle. 
For the females of such insects, moreover, a large egg mass 
is no fatal burden, and we find that the eggs are well pro- 
vided with yolk. The presence of this yolk enables the embryo 
to undergo a large part of its development within the egg. 

In the earlier Palaeozoic times none but these generalized 
insects existed. But as the struggle for existence increased, 
these insects began to adopt more fully the new environment 
which had become available to them when wings were evolved ; 
as the pressure of life increased more and more this.specializa- 
tion became more and more marked, till there was produced 
the marvellous diversity of form that exists to-day. 

Now this specialization must have had two marked con- 
sequences : — 

(a) As the insects: began to adopt a more active type of 
existence such as we see in the Diptera, Lepidoptera, and 
many Coleoptera, it became increasingly hazardous, or even 
impossible, to moult during this period of active life. It 
would be clearly impossible for a butter-fly or a blow-fly to 
undergo a moult at the present day. Moreover, the cuticle of 
these specialized insects constitutes a considerable proportion 
of the body weight. The casting and reforming of this, several 
times, would be an insurmountable strain on the insect’s 
economy. 

(b) With increase in activity it became more and more 
impossible to carry large masses of yolked eggs. Either the 


501 


quantity of yolk within the eggs would have to decrease, or 
the number of eggs would have to diminish. While the latter 
may have occurred, there is no doubt that the former process 
has predominated, till in the chalcid wasps we find eggs in 
which yolk may be almost absent. 


Under the influence of the first factor (a), with increase 
in specialization it became increasingly necessary that feeding 
should occur earlier, and ever earlier in the free living period 
of the insect. Under the influences of the second factor (0), 
it became necessary that the larva should hatch in a more 
and more incompletely developed condition. The result of 


_ the co-operation of these two processes has been very marked. 


__ aa 


In the less specialized insects as growth became more con- 
centrated in the earlier part of the free living period, the 
imaginal cells had to adapt themselves temporarily to rapid 
feeding conditions; only after a considerable time did the 
delayed differentiation occur. The metamorphosis here is very 
simple (Odonata), and the cells which have already begun to 
specialize in a certain direction (that of rapid feeding and 
growing) can, nevertheless, evidently attain the imaginal con- 
dition with comparative ease. The Coleoptera emerge from 
the egg in a more primitive condition, and while some tissues 
can still rejuvenate, others are unable to do so; they die and . 
the adult organs are formed from imaginal discs. In the 
Muscidae this process has gone much further. But in the 
chalcid wasps the divergence from the imaginal condition is 
most marked of all. It it customary to regard the Muscids 
as showing the most marked of metamorphoses, but in this 
théy are far surpassed, it seems to me, by the chalcid wasps. 
The state of embryonic development in which their larvae 
hatch has been pushed so far back that the head is still in a 
bisegmentated condition, appendages are almost entirely 
absent, malpighian tubes are not yet formed, the mandible 
may still exhibit the crustacean palp, and the proctodaeal 
ingrowth has not yet opened into the archenteron. So 
specialized, moreover, has the larva become to absorbing food 
rapidly, that it does not apparently even take time, as far as 
I could observe, to excrete nitrogen. All its food is stored 
in the form of hypertrophied larval cells within its body, and 
not till the cells attain their critical volume does transforma- 
tion occur. 


It is scarcely necessary to point out that as the insects 
gradually developed metamorphoses, the instincts of the 
parents had to undergo marked modification. As feeding had 
to become concentrated at the beginning of life, they had to 


_ deposit their ova in places where such food was to be obtained. 


= 


502 

It is necessary finally to point out the changes through 
which the tissues must have passed as the metamorphosis 
gradually evolved. As the simple ‘‘rejuvenation” meta- 
morphosis evolved,.the cells of the imago had to “‘learn’’ to 
adapt themselves to a period of rapid feeding, perhaps on 
special foods, in the early part of the life cycle. As the larval 
condition, which had then become initiated, became more 
marked greater specialization of cells resulted. For a time 
these cells were still able to rejuvenate themselves. But as 
specialization of the imago (and consequently also of the larva, 
though in a different direction) proceeded, these larval cells 
must have found it increasingly difficult and finally impossible 
to transform themselves into the imaginal cells from which 
they had phylogenetically descended. Death must eventually 
have resulted in some of the cells. Now as specialization 
increased still further more and more of these larval cells must 
have perished; in order that the individual should survive 
other larval cells would have to undergo a decrease in 
specialization. And as the present-day specialization of the 
imago was gradually attained there would thus have had to 
occur two parallel processes within the larva—an increase of 
specialization of some cells (the true larval cells) and a 
decrease in specialization, 2.e., a retention of embryonic char- 
acters in others. There would in this way be formed imaginal 
discs. That this is the phylogeny of imaginal discs is clearly 
shown by the fact that these structures are always found in 
close connection with the structure of the larva to which they 
correspond ; ¢.g., in Nasoma the antennal nerve of the imago 
arises from a cluster of cells in the brain lying very close to 
the antennal ganglion of the larva; or, to take a simpler case, 
the imaginal disc of the adult mandible lies in very close 
communication with the mandible of the larva. If the 
mandible of the larva had been evolved quite independently 
of the imaginal mandible this would not have been the case. 


To recapitulate, then, in the struggle for existence which 
has been going on among insects since Palaeozoic times, the 
possession of wings and a hard cuticle has enabled the insects 
to undergo marked specialization. Active flight made the 
carrying of numerous heavily yolked eggs impossible, and as 
the laying of numerous eggs has remained essential the quan- 
tity of yolk material has gradually diminished, reaching a 
minimum in the chalcids. On the other hand, the increased © 
loss to the animal economy sustained by the moulting of 
imaginal cuticles has necessitated that growth in size of the 
body should become more and more concentrated at earlier 
parts of the free living period; ultimately moulting has dis- 
appeared in the life of the imago. These two processes have 


SS 


e 


503 


resulted in a gradual shifting of the period of growth to the 
beginning of the free living period; and have at the same 
time forced the larvae to emerge from the egg in increasingly 


earlier periods of individual development. The processes of 


accelerated growth and of premature emergence from the egg 
have reached their maximum not among Muscids, as is usually 
supposed, but among the chalcid wasps. 

By this.gradual concentration of the growth period to the 
beginning of the post-embryonic life a new organism which 
may have to fit into a wholly new environment has been 
produced—the larva. Coenogenetic modifications, almost as 


‘interesting as the adaptations of the adult insect, have 


arisen ; but it is in the protected parasitic larvae that we must 
look for cases in which the processes of premature hatching 
and rapid feeding, unfettered by a complex environment, 
have proceeded the farthest ; and it is among the chalcid wasps 
that the most profound recovery from larval specialization— 
metamorphosis—is to be found. 

It is possible now to obtain a clearer conception of the 
significance of the pupa. When specialization first developed, 
return to the imaginal condition was doubtless by means of | 
cellular rejuvenation. Increasing specialization of the imago 
produced, automatically, as I have shown above, increased 
specialization towards rapid food absorption. When recovery 
by rejuvenation from this specialization became impossible, 
continued specialization of some larval cells must have been 
accompanied by decreased specialization of others; ultimately 
the latter would have remained as embryonic cells, and 
imaginal ‘‘discs’’ were the result. 

Now in the early stages of evolution of the larval form, 
the resulting metamorphosis must have been of a very simple 
type, differing little, indeed, from direct development; a 
simple rejuvenation of tissues, such as occurs still to a certain 
extent amongst Coleoptera, must have occurred. Nothing is 
known of the metamorphosis of the dragon-flies, but a cell 
rejuvenation metamorphosis among these forms may be pre- 


dicted with considerable confidence. There is no extensive 


tissue death, apparently, and it is not possible here to speak 
of a pupa in the usual sense of the word. But as the specializa- 
tion became more marked, extensive tissue death, and 
corresponding tissue regeneration occurring late in larval life, 
brought about an absolute break in the orderly developmental 
process, and the actual development of the less generalized 
characters of the insect did not begin till after the larval 
tissues had died. This metamorphosis begins considerably 
before the last ‘‘larval’’ moult; eventually when the larva 


_ does moult there comes to view a wholly different organism— 


504 


we can see now the imago, whose development has been so long 
delayed, and which would have appeared in this state in the 
egg if growth of the imago could have been possible. This 
‘larval’? moult in the different orders takes place when the 
embryo is in various states of development. In Lepidoptera 
the appendages are only feebly developed; the hard secretion 
of the exposed epithelium effectually serves to hide these. 
In the chalcids appendages are always well developed ; indeed, 
in this group it is possible to say that the period preceding 
the pupal moult is the period of growth in size of the external 
form of the developing imago, the pupal period the period of 
its differentiation. This generalization cannot, however, be 
applied to the internal organs; it serves merely to emphasize 
the fact that the peculiar organization of the external features 
of insects have mainly been responsible for the evolution of 
metamorphosis. It is scarcely necessary to remark, that the 
view so often held, vz., that metamorphosis commences in 
the pupal period, is quite erroneous ; for some few tissues this 
is true, but more usually the most profound changes occur 
under the shelter of the old larval cuticle. . 

It is interesting, in conclusion, to observe that the insects 
do not, after all, provide an exception to the generalization 
of Von Baer, that resemblances between different species 
become closer as we examine ever earlier stages of their 
embryonic development. It has more than once been remarked 
that the pupae of related insects are more alike than their 
larvae; but when we remember that the larva is a younger 
product of evolution than the embryo which succeeds it in 
development, and which is revealed at the pupal moult, then 
our confidence in the law, the truth of which has lately been 
so much questioned, must become stronger than ever. 


SUMMARY. 
A. External Features. 


The subject of this investigation is a small chalcid wasp, 
Nasonia, parasitic on muscid pupae, and world-wide in dis~ 
tribution. 

The larva, which is composed of fifteen segments (2 head, 
3 thoracic, 10 abdominal), feeds for three days and then begins 
to metamorphose; a day later the contents of the intestine 
are voided (defaecation period), another day later, after the 
developing imaginal discs have grown into the form of the 
imago, the ‘‘larva’’ moults ‘and discloses the pupa. Further 
development of imaginal organs and disintegration of larval 


se 
wa 
t 
s ‘A 
oY 
y 


505 


tissues occur during the pupal period. On the fourth day of 


pupal life the integument chitinises and the imago is seen 


lying within the transparent pupal sheath. 

The development of the head from the first two segments 
is fully described. The head appears to be composed of five 
primitive segments. The mouth appendages develop by the 
evagination of imaginal discs. The latter are seen in the first 
larval instar as thickenings of the integument, consisting of 
minute embryonic cells. These, like the anpendages of the 
legs, wings, and abdomer (in females) become invaginated 
during larval life as the surrounding larval cells increase in 
size, and only evaginate again at metamorphosis. These, and 
all other imaginal tissues, are to be observed in the larvae at 
all stages of their development. The most noteworthy feature 
in the mouth appendages is the occurrence of a mandibular 
palp. In the antennae, organs of smell, touch, and hearing 
are described. 

The compound ‘‘thorax’’ is described as consisting of 
three thoracic segments, as well as the first abdominal: and 
dorsal part of the second abdominal. The lower part of the 
second remains as the petiole. This is contrary to accepted 
views. Legs and wings are formed as outgrowths from 
invaginated discs on the ventral side of the thoracic segments. 

The ovipositor is formed by the outgrowth of three pairs 
of imaginal discs on the twelfth, thirteenth, and fourteenth 
segments; they come into intimate relation and co-operate to 
form the complex ovipositor. 

The penis is formed as a remarkable modification, accom- 
panied by partial invaginations, of the hinder abdominal 
Segments. 


L. Histological Changes in Intequment. 


The larval cells, having grown greatly in size during the 
feeding period, disintegrate; in part they dissolve in the 
blood; in part they are removed by leucocytes. The minute 
imaginal cells develop at their expense and no break in the 
integument is ever to be seen. The underlying somatopleure 
also metamorphoses. 

Bristles are formed as partial chitinisations of cells or 
groups of cells which have grown out into the form of bristles. 
Pubescences are formed as chitinisations of the frayed exteriors 


of other cells. Phragmas are formed as cleft-like invaginations 


of the integument; false phragmas by the downgrowth of the 


Margin of segments. 


The eye is represented in the first intsar by an integu- 


- mental thickening consisting of three layers of cells. From 
the outer develop the lens, and vitreous cells; from the middle 


506 


the ‘‘retinula’’ and pigment cells; from the lower the rhab- 
dome cells. The cells arrange themselves in little groups 
(ommatidia) each consisting of one central rhabdome cell, 
surrounded by seven sheath (‘‘retinula’’) cells (of which one 
later disappears), which are, in turn, surrounded by four 
pigment cells. At the outer end are formed four vitreous and 
two lens cells. The latter surround the former*and their 
upper ends secrete the lens. The rhabdome cell chitinises. 
An ingrowth from the ectoderm which surrounds the eye forms 
a membrane beneath the developing eye (‘‘perioptic mem- 
brane’’). Its cells serve as neurolemmae for the cells of the 
optic ganglion. The remarkable process of the development 
of the eye innervation in connection with the perioptic mem- 
brane is fully described. | 

The ocelli are similarly modifications of the integument 
only, and the nerve reaches them, quite independently, from 
the brain. 


C. Resmratory System. 

The larva has a pair of longitudinal tracheal trunks, 
connected by transverse trunks in front and behind. The 
longitudinal trunks open to the exterior by four short stig- 
matic trunks, which. increase to nine in the second larval 
instar. The air is carried to the tissues by extraordinary 
tracheoles, of a kind not hitherto, apparently, described. Hach 
branching system, of which there are two to five in a segment, 
is simply a highly branched hollow cell, formed from a greatly 
‘modified cell of the tracheal epithelium (giant tracheoloblast). 
Increase of complexity during larval life is produced by 
growth of the size of these cells, and by further branching. 

The whole tracheal system degenerates at metamorphosis, 
partly dissolving in the blood, partly removed by leucocytes 
after degenerating. A simultaneous regeneration from 
embryonic cells which form imaginal ‘‘nests’’ at the bases of 
all the stigmatic trunks, prevents any discontinuity in the 
tracheal system occurring. The embryonic cells grow over the 
dead larval cells of the main trunks; growing out in places 
they now form the true branching multicellular tracheal 
vessels in the head, alitrunk, and abdomen. Some of them 
are modified into the great thoracic air sacs. The “‘spiral’’ 
intima of the tracheae is formed as a chitinisation of ridges 
formed on the inside of the cells which compose the tracheal 
epithelium. 

D. The Muscular System. 

The larval muscular system begins to degenerate in places 
before defaecation, ¢.g., in the thorax, embryonic cells 
(myoblasts) begin to crawl over certain dorsal longitudinal 


i 


507 


thoracic muscles which are beginning to lose their striations 
and ultimately form the wing muscles. They penetrate the 
muscles, which soon become riddled with these cells and 
ultimately absorb them; in their place is formed a pair of 
bands of myoblasts. Some of these myoblasts fuse in five 
longitudinal columns within each band, and the syncytium 
remains as the sarcoplasm of the future ‘‘wing muscles.” 
Other myoblasts send off processes into these columns and 
form the sarcostyles of Schafer, which are therefore fibres, not 
fibrils, as usually supposed. Each band then breaks up into 
its five constituents, and the great ‘‘wing’’ muscles of the 
thorax are formed. The ‘‘muscle insertions” are always 


integumental cells. | 


a 


Other muscles of the larva, which may show all kinds of 
disintegration processes, may become absorbed by leucocytes ; 
others, again, dissolve slowly in the blood stream, usually 
throwing out large rounded globules as they doso. A specially 
remarkable case of disintegration is seen when the whole of 
the minute sarcous elements are cast out as a fine shower of 
“bacillus-like’’ rods into the blood stream where they dissolve 
(fig. 104). 

In all cases the adult muscles are regenerated from 
embryonic cells (myoblasts), like leucocytes in appearance, 
which have lain dormant during larval life. They unite one 
after the other to form syncytial columns, one or more cells in 
thickness. If more than one ceil in thickness the columns 
may (head, leg, and ovipositor muscles) or may not (pharyn- 
geal dilators) become pulled apart to form a number of 
narrower columns. Each of these columns undergoes fibrilla- 


tion and striation by a method quite different from that 


described in the ‘‘wing’’ muscles. In structure the muscles 
always present striations in the form of double spirals. Certain 
unicellular intestinal muscles may be markedly branched, each 
branch consisting of only a few fibrials. Striation is here 
truly transverse. 


E. The Intestine. 


The larval intestine consists of fore-, mid-, and hindguts. 
The latter does not open into the midgut till towards the end 
of larval life. This marks the defaecation period. 


The imaginal tissues are (1) a ring around the posterior 


_ end of the oesophagus, (2) scattered cells at the bases of the 
larval cells of the midgut, (3) cells surrounding the anterior 
_ parts of the rectum. Salivary glands are well developed ; 
into the midgut open three hepatic caeca; malpighian tubes 
are absent in the larva. 


P2 


4 
508 7 


On the fourth day of larval life the foregut and rectum 
degenerate and are rapidly regenerated by cells growing from 
the imaginal rings. The oesophagus is also partly regenerated 
from the head integument, which creeps in through the mouth. 
The cells of the midgut disintegrate by a remarkable process 
of globular degeneration and fall into the lumen of the gut; 
the epithelium is rapidly regenerated by the ‘‘imaginal’” 
cells of the midgut. The hepatic caeca become bodily drawn 
in through the walls of the degenerating larval midgut. This 
is produced by pressure from the regenerating epithelium. 
The function of this epithelium is to absorb the disintegrated 
cells. The anterior portion of the epithelium then itself 
breaks up into a fine débris and is absorbed by leucocytes ; 
the posterior part remains as the stomach. Meanwhile the 
imaginal ring of the oesophagus has formed a great cone of 
cells, which temporarily has closed the midgut in front. The 
cells of this cone now grow back through the thorax and fuse 
with the stomach; they differentiate to form gizzard and crop. 
The malpighian tubes grow out from the anterior part of the 
hindgut in the defaecating larva. The hindgut bends upon 
itself on account of rapid cell proliferation ; the anterior part 
is the small intestine, the hinder the short rounded rectum. 
Within its walls is formed a pair of rectal glands, by thicken- 
ing of the epithelium. | 

The salivary glands, after disintegrating, are phago- 
cytised. A single salivary gland is formed by ingrowth of 
cells from the regenerated oesophageal epithelium. Only one 
salivary gland occurs in the imago. 


F. The Ductless Glands. 

(1) The Oewocytes.—The larval oenocytes grow in size 
but do not proliferate; at metamorphosis they simply dis- 
integrate. Leucocytes may at times aid in their removal. The 
imaginal cenocytes are formed from small clusters of cells 
which grew inwards from the ectoderm in the early larva. 
They separate and scatter themselves between the fat cells. 

(2) The Lateral Intestinal Glands are a pair of long 
chains of glandular cells lying just beneath their lateral 
hepatic caeca. They disintegrate during metamorphosis. 


(3) The Dorsal Abdominal Glands.—The imaginal anlage 
of these is a band of cells lying in the young larva dorsally 
on the end of the midgut. They grow and proliferate late 
in larval life and resemble empty fat cells. During late pupal 
life they assume a glandular appearance. They persist through- 
out imaginal life and are not pericardial cells or young fat 
cells. 


509 


G. The Fat-hody. 

The cells of the fat-body do not proliferate during larval 
life. They grow in size and store fat and other (protein /) 
foodstuffs in their cytoplasmic meshwork. Before defaecation 
excretory crystals may accumulate within them, though these 
soon disappear when the malpighiaw tubes form, and are cast 
into the stomach, where they accumulate during the pupal 
period. 

The storage products gradually disappear during pupa- 
tion, as the imaginal tissues grow at their expense. Fre- 
quently the remnants of the fat cells, deprived of their storage 
substances, are phagocytised. Others persist throughout pupal 
and imaginal life. There is no regeneration of fat-body. 
Some of the fat cells are seen to have a capacity for limited 
- phagocytosis. 


H. The Gonads. . 

These occur in the youngest larvae as a pair of club- 
shaped masses attached to the ventral body wall. They 
begin to grow at metamorphosis and develop directly into the 
adult organs. The male organs often show ripe sperms 
already in the third day of pupal life. The ovaries continue 
to develop during imaginal life. 


IT. The Nervous System. 


In the larva there is a brain above the oesophagus con- 
nected to a ventral nerve cord consisting of twelve ganglia, 
the last of which consists of three fused ganglia. A single 
. Stomotogastric ganglion is present. The nerve cord and brain 
are composed of larval cells and imaginal neuroblasts. Dur- 
ing metamorphosis the former degenerate, and the latter 
proliferate. In the nerve cord the dead cells which form 
masses of necrotic tissue there, are quickly absorbed: into 
the degenerate strands of fibres that run along the cord, the 
new nerve cells send their growing nerve fibres, and no dis- 
continuity is to be observed. In the peripheral nerves, the 
fibres break into globules which, passing out, dissolve in the 
blood or are phagocytised. All these changes take place 
within the splanchnopleural covering of the nerve cord, which 
itself metamorphoses by imaginal cells replacing dead larval 
eells, and no discontinuity occurs in it. It follows, therefore, 
that no discontinuity is to be observed in the nerve cord and . 
peripheral nerves as a whole, although the most profound 
changes are taking place within it. These changes are com- 
pleted before pupation; a migration of nerve cells (7.¢., of 
ganglia) then commences and the concentrated nervous system 
of the imago is formed. 


510 


Similar changes occur in the brain ; but within it the dead 
larval elements lie for more than a day as masses of degenerate 
cells, before these are absorbed by the imaginal cells. The 
first ventral ganglion is merged into the brain, and it is much 
more complex than in the larva. 

Leucocytes do not take part in the metamorphosis. 


J. The Vascular System. 

The blood acts as the medium into which the nutrient 
degeneration products of the larva are poured and from which: 
the imaginal cells again take them. Often the leucocytes, 
which proliferate considerably during metamorphosis, may 
aid in the removal of the dead larval elements by phagocy- 
tising them. 

The larval heart consists of a long tube, provided with 
ostia, and lying within a delicate pericardium. Imaginal 
cells lie within the heart walls and regenerate it at the time 
of defaecation. The larval pericardium undergoes total 
degeneration. In its place is formed a band of cells right. 
along the ventral side of the heart; the cells originate from 
an imaginal disc lying ventral to the heart, just above the 
end of the hindgut. This band of cells then grows upwards. 
and completely surrounds the heart, forming a two-layered 
organ, which becomes muscular behind (heart), remaining 
non-contractile in front (aorta). There is therefore no true 
pericardium in the imago. 

K. The Insect Metamorphosis. 

Insect metamorphosis is brought about by an extensive 
tissue death, due to the hypertrophy of the larval cells. 
Death of the cells is due to an automatic starvation, due to 
the fact that the cell contents, which increase as the cube 
of the radius of the growing cells cannot be nourished 
indefinitely through the cell membrane, whose area increases 
only as the square of the radius. The hypertrophy must also 
cause disorganization, as the distances through which diffu- 
sions, etc., within the cells have to take place, become 
appreciable. 

Metamorphosis has been evolved as an outcome of 
specialization by the imago. A hard thick cuticle has 
necessitated concentration of the growth period to the begin- 
ning of the free living period, and the active life led by the 
adults has made the carrying of large quantities of heavily 
yolked eggs impossible. With continued decrease in the 
quantity of yolk, the larva has had to emerge at ever earlier 
stages of its development. At metamorphosis we see the con- 
tinuation of the interrupted development, and a recovery of 
the organism from larval specialization. 


511 


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Samson, K. 
1908—Ueber das Verhalten der Vasa Malpighii und die 
excretorische Funktion der Fettzellen wahrend 
der Metamorphose von Heterogenea limacodes. 
Zool. Jahrb. Anat., Bd. 26, p. 403. 
SAVIGNY. 
1816—Mémoires sur les Animaux sans Vertébrés, Pt. 1. 
Paris. 
ScHarer, E. A. 
1891—On the Minute Structure of the Muscle-columns, 
or Sarcostyles, which form the Wing-muscles of 
Insects. Proc. Roy. Soc., London, vol. 49, 


p. 280. 
SHarp, D. 
1895—Insects. Cambridge Natural History. 
SupPino, F. 


1900—Osservazione sopra fenomeni che avvengono durante 
lo sviluppa postembrionale della Calhphora 
erythrocephala. Bol. Soc. Entom. ital., Anno 
32, p. 192. Also, Atti della R. Acad. dez 
Lincei, 5, 9. 
TILLYARD, R. J. 
The Biology of Dragon-flies. Cambridge, 1917. 


515 
"TOWER. 


1909—The Origin and Development of.the Wings of 
Coleoptera. Zool. Jahrb., vol. 17. 
Vaney, C. 


1902—Contributions 4 lVétude des larves et des méta- 
morphoses des Diptéres. Ann. Univ. Lyon., 
new series, fasc. 9. 


Van Rees, J. 


1884—Over intra-cellulaire spijsvertering en over de 
beteekensis der vitte bloedlichaampjes (Maanblad 
voor Natuurvetenschappen IT.). 

1889—Beitrage zur Kenntniss der inneren Metamorphose 
von Musca vomtaria. Zool. Jahrb. Anat., 5, 3. 

VeERsON, E. 

1898—Zur Entwicklung des Verdauungscanals beim 
Seidenspinner. Zool. Anz., 21. 

1905—Zur Entwicklung des’ Verdauungscanals bei 
Bombyx mori. Zeit. f. Wiss. Zool., vol. 82, 
p. 523. 


VIALLANES, H. 


1882—Recherches sur Uhistologie des Tusectes Ann. Sc. 
Nat. Zool., (6), 5, 14. 

1884—-Etudes histologiques et organologique sur les 
centres nerveux et les organes des sens des 
animaux articules. 2e Mémoire. Le ganglion 
optique de la Libellule. Ann. Sc. Nat. Zool., 
(G):55 18. | 

1885—3e Mémoire. Le ganglion optique de quelques 
larves de Dipteres. Ann. Sc. Nat.- Zool., 
(Gy; 54, 19. 

1887—4e Mémoire. Le cerveau de la quépe (Vespa crabro 
et V. vulgaris). Ann. Sc. Nat. Zool., (7), 5, 2 

WEISMANN, A. 

1864—Die nachembryonale Entwicklung der Musciden. 
Zeit. f. Wiss. Zool., Bd. 14. 

1866—Die Metamorphose der Corethra plumicorms. Zeit. 
f. Wiss. Zool., Bd. 16. 

“WEISSENBERG. 

1907—Ueber die Oenocyten von T'orymus nigricorms, 
mit besonderer Beriicksichtigung der Meta- 
morphose. Zool. Jahrb. Anat., Bd. 23. 

WHEELER, W. R. 

1890—Neuroblasts of Arthropod embryos. Journ. 
Morphol., 5, 4. f 

‘WIELOWIESSKI, H. 

1886—Uber das Blutgewebe der Insecten. 


516 


Reference to Letiering. 


a.: Antenna. ab.l-ab.10: Abdominal segments. a.d.: 
Imaginal] disc of antenna. a.g.: Accessory glands. a.g.i.: Im- 
aginal antennal ganglion. a.g.J.: Larval antennal ganglion. 


a.l.: Antennal lobe. an.: Anus. a.u.: Antennal nerve. ant.: 
Second (true) antenna. a.p.: Adhesive pad. app.ab.10: Append- 
age of tenth abdominal segment. a.s.: Antennal nerve strand. 
b.m.: Basement membrane. br.: Brain. c.: ‘Drum-shaped’’ 
chamber. ¢.0.c. Gircumoesophageal connective. cr. : hee 
dai: Dorsal apaenen giand. d.g.: Degeneration globules. 
e.: Hye. e.op.: Epiopticon. ex.m.: Heenan muscle. f.: Sarco- 
style. f.b.: Fat-body. 9 f.c.: Follicie cells. ‘f.2.: > Wore eee 
fl.m.: Flexor muscle. g.l-g 12: Ganglia of ventral nerve cord. 
gl.: Lubricating gland. gz.: Gizzard. h.: Heart. h.c.: Hepatic 
caecum. i.c.: Intestinal calls ie ee Imazinal dise of integument. 
i.i.c.: Imaginal integumentary cells im.c.: Imaginal cell. 
imt-C,, Integumentary (larval) cell. i.o.c.: Imaginal oesophageal 
cell. i.oen. Imaginal Lie: bess 10,0. : Imaginal tracheal cell. 
rs Hs ie Imaginal tracnea. Jaw of larva. k.m.: Krause’s mem- 
brane. 1.: Leucocyte. ee Legs. lab.: Labium.  Ibr.: 

Labrum. l.c.: Lens cells. I.d.: Imaginal* disc of leg. Li.g. = 


Lateral intestinal gland. 1.m.: Longitudinal muscle.  1.0.c.: 
Larval oesophageal cell. 1.p.: Labial palp. Is.: Lens. Ls.: 
Larval cuticular sheath. l.t.c.: Larval tracheal cell. I.v-e.; 


Larval cell. m.: muscle. malp.t.: Malpighian tubule. md.: 
Mandible of imago. m.d.: Imaginal disc of mandible. md.p.: 
Mandibular palp. m.g.: Mid gut. m.i.: Muscle insertion. 
ms.tx.: Mesothorax. mth.: Mouth. mt.tx.: Metathorax. mx.: 
Maxilla. mx.p.: Maxillary palp. myb.: Myoblasts. n.: Nucleus. 
n.c.: Nerve cell. ne.: Nutritive cell. ng.c.: Neuroglia cell. 
nml.: Neurolemmal cell. u.0.: Nerve to oral appendages. nrb.: 
Neuroblast. n.s.: Nerve strand. n.t.: Dead larval tissue. n.v.: 
Nerve. o.: Ovum. oc.: Ocellus. od.: Oviduct. oen.: Oenocyte. 
oes.: Oesophagus. o.g.: Optic ganglion. o.g.2: Middle optic 
ganglion. o.g.3: Inner optic ganglion. o.gl.: Ocellar ganglion. 
om. : Ommatidium. o.m.: Oblique muscle. o.n.s.: Ocellar nerve 
strand. o.p.: Opticon. op.n.: Optic nerve. ost.: Ostia. ov.: 
Ovary. ov ip. : Ovipositor. ovip.l-ovip.3: Imaginal dises of ovi- 
positor, or the structures into which they have developed. p.b.: 

Polar body. p.c.: Pigment cell. p.d‘m.: Pharyngeal dilator 
muscle, ped.: Beceeedinies, ph.: gharyit phr.: Phragma. 
p.m.: Perioptic membrane. p.m.c.: One of the remarkable 
branching cells of the perioptic membrane. p.op.: Periopticon. 
pt.: Petiole. pr.tx.: Prothorax. r.d.: Imaginal disc of rectum. 
rect.: Rectum. rect. g. : Rectal gland. rh.: Rhabdome (or 
rhabdome cell). rp.c.: Replacing cell of midgut epithelium, 7.e., 
the anlage of the tempor ary pupal midgut. s.: Syncytial columns 
of wing muscles. sal.d.: Salivary duct. sal.g.: Salivary gland. 
s.a.o.: Sensory appendage of ovipositor. s.c.: Sheath cell (‘‘reti- 
nula”’ cell). s.i.: Small intestine. sp. Splanchnopleure. 
sp.l-sp.10: Lateral spiracles. — spl.: Mesodermal somatopleure. 
sp.n.: Nucleus of a splanchnopleure cell. 8.7. : Sensory rod. st.: 

Stomach. stg.: Stomatogastric ganglion. “t.: Tendon. t.: 
Testis. t.o.: Ovarian tubules. trb.: Developing tracheoloblast. 
trl.: Tracheole. 1 Oe Longitudinal tracheal trunk. t.t.t. : 

Transverse tracheal trunk. v.: Vagina.  v.e. Vitreous cell. 
v.d.: Vas deferens. v.n.c.: Ventral nerve cord. v.s.: Vesicula 


517 


seminalis. vsc.: Vesicle. w.l-w.2: Wings. w.d.: Imaginal disc 
of wing. W.1. ne insertion. w.m.: ‘‘Wing’’ muscles. w.m.b.: 
“Wing’’ muscle band. x.: Various structures referred to in text. 


; DESCRIPTION OF PLATES. 

(a 

‘ PratEe XV. 

My Fig. 1. Larva of Nasonia towards end of first instar, in 
ventral view. The muscles are shown on one side; on the other 


are seen the respiratory vessels. The nerve cord is also visible 
|i Fig. 2. Larva at end of feeding period (x55), showing the 
_ fifteen imaginal discs of the integument (heavily shaded), one 
_ (pair) in each segment. The respiratory system is also seen; the 
__ alimentary canal is shown in outline. 
7 Fig. 3. Anterior end of larva at time of defaecation (x 150). 
Metamorphosis has commenced, and the various appendages are 
clearly seen through the transparent larval sheath. Note the 
| fat-body showing up through the integument; note also the 
| mandibular palp. 

Fig. 4. Abdomen of imago, in ventral view. The numbers 
refer to the abdominal segments. 
. | Fig. 5. Integumental imaginal dise of larva shown in fig. 2 
(180). showi ing the invaginated leg anlage. 
. Fig. 6. Anlage of leg, from the same specimen, viewed in 
' optical section (x 300). Note the mesoderm extending into the 
| hollow appendage. 
| ; PuateE XVI. 


4 Fig. 7. Fresh pupa (x50), showing migration of first two 
abdominal segments. 

Ra Fig. 8. The same, four days later (x 50). Note the exten- 
_ sive shrinking that has occurred. Note also final position of 
_ migrated abdominal segments. 

i Fig. 9. View of propodeum and adjacent segments of pupa 
bs two days old (x70). Note the migrated abdominal segments ; 
also the complex wing insertions. 

*: Fig. 10. Transverse section through first thoracic segment 


| of larva in first instar (x1000). The complete section has not been 
drawn. Notice the imaginal elements in the integument, nerve 
- eord, and intestine. 
Fig e. ll. Antennae of female wasp, imago (x120). The 
bristles are not shown. Note the auditory organs. 
' Fig. 12. Head of metamorphosing larva, about 17 hours 
before pupation, 7.e., seven hours later than fig. 2 (x 150). 


Prate XVII. 


} Fig. 13. Head of mature larva at cessation of feeding, 

viewed ventrally, and slightly from one side, showing all the 

imaginal discs of ‘the mouth appendages (x159). Note the double 
_ nature of the labrum; also the true antenna (ant.), which later 
_ disappears. Mandibular palps are not visible with certainty. 

a Fig. 14. Mouth appendages of pupa aged two days (x150). 
_ Note shrinking within the pupal sheath. 

Fig. 15. Longitudinal section of part of the antenna of 

i erty six hour pupa (x 450). 


4 

Fig. 16. Leg of pupa two days old (x90). The leg muscles 
are clearly seen. 

Fig. 17. Trochanter of leg of imago (x500) to show the 
nature of the muscle insertions. 

Fig. 18. Insertion of tendons of femoral muscles on tibia 
(x 500). 

Fig. 19. Tactile bristles on first tarsal segment of first leg, 
imago (x1500). Note the curious modification of the somato- 
pleural mesoderm cells to act as neurolemmae for the nerve fibres. 

Fig. 20. Developing ovipositor from larva seven hours after 
AER ction (x 150). 

Fig. 21. The same, several hours after pupation (x180). 

Fig. 22. Ovipositor (extruded) of imago. Note the great 
musculature. 

Fig. 23.. Section through a part of the propa (thirty- 
six hour pupa) showing the powerful chitin layer (x 500). 

Fig. 24. Developing pubescence on labium, fifty-six hour 
pupa (1500). Chitinisation has just commenced. Note cell 
nuclei. 

Fig. 25. Vertical longitudinal section through posterior end 
of male, to show the migration of segments to form the penis. 
Note cell proliferation in ninth segment (x150). Fresh pupa. 

Fig. 26. Ventral view of extremity of male—two-day pupa 


(x 150). 
Prare XVIII. 


Fig. 27. Longitudinal section of posterior end of male— 
fifty-six hour pupa (x180). The cuticular sheath represents the 
position of the segments before invagination occurred, . 

Fig. 28. Integument undergoing metamorphosis, ventral 
abdominal region of larva eight hours after defaecation. Note 
the greatly hypertrophied larval cells with large nucleoli, all 
degenerating. Only a few leucocytes are present. Note the small 
onbrane cells growing over them (x 1200). 

29. Surface view of a proliferating imaginal disc. The 
larval alle are degenerating. Some of the embryonic imaginal 
cells are undergoing amoeboid movement (x), From the larva, 
shortly after defaecation (x 1000). 

Fig. 30. Section through integument of larva eight hours 
before pupation (x1200). Note the empty cell membranes (z) and 
granular degeneration globules, free, or still within cells. { 

Fig. 31. Integument from posterior extremity of eight-hour 
pupa, showing accumulation of degeneration globules amongst the 
cells of the renovated integument. A leucocyte and a degenerate | 
larval cell are also seen (x 1200). ; 


518 


™~ 


Fig. 32. Section through mid-dorsal abdominal integument. 
A disintegrated integumental cell is seen, apparently preventing 
the closing in of the imaginal cells. The heart is also seo | 
(x1200). From the larva twelve hours after defaecation. | 

Fig. 33. Integument of head showing development of scale- _ 
like ‘‘bosses.’? Note that a protoplasmic ‘‘mould’’ precedes the 
secretion of the similarly shaped cuticle (x 1500) —fift ty-two hour 
pupa. 

Fig. 34. Developing spine of leg (x1200)—fifty-two hour 
pupa. iff 
Fig. 35. Developing abdominal bristles (x 1200)—fifty-two 
hour pupa. Note the formation of a ‘“‘mould’’ by the integu-— 
mental cells, previous to cuticle secretion. 


519 


Fig. 36. Longitudinal section of antenna (x450). Defaeca- 
ting larva. Note invagination of ‘‘padding” tissues, for example 
at x. 

Fig. 37. Portion of wing of four-day pupa, showing hairs, 
and the characteristic folding. Degenerate nuclei also seen 
(x 750)... : : ‘ 

Fig. 38. Hooks of hind wing of imago, showing scattered 
wing nuclei; hook -‘‘roots’—remnants of embryonic cells—are 


clearly seen. 
Prate XIX. 


x 500). 
Fig. 42. Section through free part of ovipositor of four-day 
pupa (x600). 

Fig. 43. Vertical section through a cephalic phragma (thirty- 
six hour pupa); developing antennal muscles are seen; also a fat- 
body and degenerating leucocytes (x 1200). 

Fig. 44. Fore wing of twenty-four hour pupa. Note shrink- 
ing within the old cuticle. Note also the two clear channels each 
containing tracheoles (x80). 

Fig. 45. Vertical section through the wing of four-hour 
pupa, to show the band of mesodermal cells (k) growing down- 
wards, to enter the anterior clear channel (500). 

Fig. 46. Optical section of tip of tarsus of imago (x 750). 

Fig. 47. Section through head of larva in first instar, right 
half only shown (x 500). 

Fig. 48. Section through mandibles of larva twelve hours 
after defaecation. 

Fig. 49. Section along part of first segment of antenna, to 
show nerve endings or tactile bristles (x 1200)—four and a half day 
pupa. 

Fig. 50. A tactile bristle from ovipositor appendage, showing 
the receptor cell (x 1500). 

Fig. 51. Sense cells below cuticle of antenna] joint of imago 
(x 1500). 

Fig. 52. Longitudinal section through the ‘‘olfactory seg- 
” of antenna, showing the suspended sense organs (1200). 
Fig. 53. Surface view of one of the more anterior antennal 
segments, showing four auditory organs (x1200). 

Fig. 54. Cells of the optic imaginal dise of larva in first 
instar (x 2000). 

PLATE XX. 


Fig. 55. Vertical section through imaginal disc of eye in 
larva of first instar (x1000). 
Fig. 56. Ommatidia from developing eye at time of defae- 
cation (x 1500). 
Fig. 57. The same, eight hours later (x 1500). 
Fig. 58, a, b, c. The same, eight hours later than fig. 57 
(x1500). d. The same, in tranverse section. Note the seven 
- sheath cells (x 1500). 
Fig. 59. Ommatidium from pupa, four hours old (x1500). 
Fig. 60. Ommatidium of four-hour pupa in transverse sec- 
tion. Note only six sheath cells. 


men 


520 / 


Fig. 61. Ommptidium of pupa twenty-four Re old (x 1500). 
Fig. 62. The*same, thirty-six hour pupa (x 1500). 
Fig. 62a. The same, outer end (x 1500). 
Fig. 63. Outer part of ommatidium of fifty-two hour pupa 
showing pigmentation of sheath cells (x 1500). 
Fig. 64. Ommatidium from eye of four and a half day pupa 


Fig. 644. The same, outer end (x 1500). 

Fig. 648. Pigment cell from the same (x 1500). 

Fig. 64c. Section through vitreous cells of same (x 1500). 

Fig. 64p. Section through terminations of several omma- 
tidia at same stage (x1500). 

Fig. 65. Fresh pupa, anterior half, to show tracheal vessels 
(x 40). 

Fig. 66. Section of ocellus in two-celled stage, from the larva 
in first instar (x 1000). 

Fig. 67. Section through developing ocellus from the fresh 
pupa (1000). 

Fig. 68. The same, twelve hours later (x 1000). 

Fig. 69. The same, from thirty-six hour pupa (x 1000). 


Pratt XXI. 
Fig. 70. The same, three-day pupa (x 1000). 
Fig. 71. Vertical section through median ocellus of imago 


Fig. 72. A single sensory cell from the same (x 2300). 

Fig. 73. Pigment cell from fifty-two hour pupa. 

Fig. 74. Vertical section through the eye and perioptic mem- 
brane, es larva about to pupate (x1200). Several cells of the 
perioptic membrane are shown. The three types of processes are 
clearly seen, especially the fibrous processes, which penetrate into 
the brain. Into these the nerve cells are seen migrating, and 
one, on the extreme left, has communicated with an ommatidium. 

Fig. 75. The same, four hours later, showing nerve cells 
adhering to the ‘“neurolemmal” cells of the perloptic membrane 
(x 2000). 

Fig. 76. A simple respiratory cell from the abdomen of 
larva eight hours before pupation. An oenocyte is also seen, pre- 
senting vacuoles, evidently a sign of degeneration (x500). 

Fig. 77. Vertical section through right side of head of larva 
eight hours after defaecation (x 230). 


Pratt XXII. 

Fig. 78. Section through left side of head of defaecating 
larva (x 230). 

Fig. 79. Vertical section through developing eye of thirty- 
six hour pupa (x 400). 

Fig. 80. Vertical eeprien through right side of head of four 
and a half day pupa (x230 

Fig. 81. Central (nuclear) region of giant respiratory cell, 
from larva sixteen hours after defaecation (x 1000). 

Fig. 82. Tracheoloblast growing out from regenerating 
tracheal trunk—defaecating larva (x 1000). 

Fig. 83. Epithelium of longitudinal tracheal vessel under- 


going histolysis (x1000)._ Leucocytes are attacking the dis- 
integrating cells. 


Rp te: 


52] 


Fig. 84. Metamorphosing anterior transverse tracheal trunk, 
eight hours after defaecation (x500). Note the tracheoblasts 
advancing from the sides, beneath the degenerate epithelium, 
which has not yet begun to disappear. 

Fig. 85. Part ‘of the same vessel, eight hours later. A 
remnant of the larval epithelium is still seen. Note the larval 
spiral intima within the developing imaginal intima. Note also 
the ‘‘ridging’’ of the epithelial cells (x 500). 

Fig. 86. Metamorphosing abdominal longitudinal tracheal 
trunk, fone the defaecating larva (x1000). Note the imaginal 
cells advancing upon the degenerate larval cells, which are partly 
disintegrating, partly still intact. Leucocytes. are present, and 
three are attacking a tracheole. A large imaginal tracheole is 
beginning to grow “out from the regenerated epithelium. 

Wigs. 87. “Central nuclear region of the great tracheal cells 
undergoing phagocytic histolysis—fresh pupa (x 1000) (cf. fig. 81). 

Wig. 88. From the large regenerated tracheal vessel (i.tr.) a 
multicellular tracheal vessel is extending downwards into the 
wing. Beside it a dead larval tracheole (trl.) is being over- 
whelmed by leucocytes. The figure also shows disintegrating 
salivary gland tissue, being attacked by leucocytes. Some of the 
gland tissue has been ruptured by the growing tracheal vessel. 
(Figure drawn inverted; x800.) From four-hour pupa. 

Fig. 89. Cell proliferation of prothoracic stigmatic trunk 
(x 525). Defaecating larva. 


,Puate XXIII. 

Fig. 90. Larval tracheoles undergoing phagocytosis (x 1000) 
—four-hour pupa. 

Fig. 91. From the regenerated neck tracheal vessel (i.tr.) 
a very large tracheal trunk has grown downwards and termin- 
ated near the mouth. A larval tracheole (trl.) is degenerating, 
without intervention by phagocytes. From the base of the head 
a column of myoblasts—-developing into the head musculature— 
has grown up supporting itself upon the dead tracheole. The 
break of the column is due to its bending out of the plane of 
section. The figure may be regarded as continuous with fig. 88 
on its Jeft (four-hour pupa; x 500). 

Fig. 92. Longitudinal section of integument of defaecating 


_ larva in region of an abdominal stigmatic trunk, which will not 


be reformed in the pupa. Notice the hypertrophied larval cells 
and the proliferating embryonic cells, especially at base of trunk. 
A fat-body and a group of imaginal oenocytes are also seen 
(x 800). 

Fig. 93. Stigmatic fails of larva in first instar showing 
aL *‘nest”’ (i.t.e.). 

Fig. 94. Developing dorso-lateral air sac—fresh pupa 
(x 1000), 

Fig. 95. Portion of wall of a mature air sac (four and a half 
day pupa). Notice that a tracheole has grown out from it. Four 


nuclei, and “‘spirals’’ are also seen (x 1000). 


Fig. 96. A cell of the fat-body fr om thorax, four and a half 


=—-day pupa. Note the disappearance of storage products (x 800). 


Fig. 97. The same, a little later, being attacked by three 
leucocytes (x 1000). 

Fig. 98. A fat cell, drawn out and compressed between the 
great thoracic muscles ; ‘note diminution in quantity of storage 
material (x 1000). 


522 


Fig. 99. Portion of body musculature of larva in first instar ; 
viewed in the living larva through the transparent integument. 

Fig. 100. A single longitudinal muscle (mucle fibre) from the 
same, composed of four distinct cells. The insertion of the muscle 
on an integumentary cell is also seen (x1000). 

Fig. 101. Piece of a muscle from adult larva, showing double 
spiral striations (x 900). 

Fig. 102, Muscle nuclei: (a) from adult larva; (b) from 
larva in first instar. Note only a slight increase in nuclear size. 
The older nucleus shows three small nucleoli; there has apparently 
been no increase in the quantity of chromatin (x 1500). 

Fig. 108. A tracheole from developing forewing of larva 
eight Bois after defaecation. 


PrateE XXIV. 


Fig. 104. Part of degenerating oblique abdominal muscle 
(fibre) Meni a disorganization of the sarcomeres—defaecating 
larva (x1200 

Fig. 105. A degenerated ventral longitudinal abdominal 
muscle (fibre) being attacked by leucocytes. Note the degenerate 
nuclei, fresh pupa (x1200). 

Fig. 106. Dorsal longitudinal see into which myoblasts 
have penetrated, fresh pupa (x1000 

Nie... 107... The same“ Cx a 200). Amoeboid myoblasts have 
entered the extruded sarcoplasm. 

Figs. 108, 109. The same (x 1000) in various degrees of meta- 
morphosis. 

Fig. 110. Degenerating circular (oblique) muscle from pro- 
podeum. Myoblasts are extending over the degenerate fragments 
of larval muscle. A few leucocytes and extruded degeneration 
globules are also seen (1200). 

Fig. 111. A regenerated abdominal muscle, thirty-six hour 
pupa. Cell limits are still indistinctly visible. Fibrillation has 
not yet commenced (x 1200). 

Fig. 112. Longitudinal section through proliferating myo- 
blasts of mesothorax, to form the great ‘‘wing’’ muscles (x 900) 
—defaecating larva. 

Fig. 118. The’same, in prothorax ; iaealnwee extending over 
a larval muscle, in which striations are still visible—defaecating 
larva (x 1200). 

Fig. 114. The great column of myoblasts in head, being 
drawn apart into its constituent muscles (x800)—thirty-six hour 
pupa. The column, at a much earlier stage, is shown in fig. 91. 

Fig. 115. <A small portion of the same, to show traces of 
the separate myoblasts (1500). 

Fig. 116. The same, fully developed—four and half day 


pupa (x1200). 
Puats XXV. 


Fig. 117. Longitudinal section through the head of defaecating 
larva near mid-line (x500). The oesophagus, salivary duct, 
pharyngeal dilator muscles and integument are undergoing 
metamorphosis. 

Figs. 118, 119. A pharyngeal dilator muscle degenerating 
without the intervention of leucocytes—defaecating larva (1200). 

Figs. 120, 121. Myoblasts overwhelming pharyngeal dilator 
muscles. In fig. 120 the striations are still present. 


523 


Fig. 122. Regenerated pharyngeal dilator muscle—ifresh 


pupa (1200). 


Fig. 123. The same; fibrils present; striations are already 
plainly visible below, still developing above. 

Fig. 124. The same—four and a half day pupa. This muscle 
shows several ‘‘roots,’ which is unusual. The structure of the 
pharynx is also shown. Note the thick muscle layer (x1200). 

Fig. 125. A branched contractile cell from the crop—four 
and a half day pupa (1200). 

Fig. 126. Longitudinal section through the insertions of 
adjacent body muscles (x800)—adult larva. 

Fig. 127. Longitudinal section of insertion of a larval muscle 
on the integument (slightly diagrammatic). The spindle-shaped 
sarcomeres are clearly seen. Also Krause’s ‘‘membrane’’ (k.m.) 
(x 1500). 

Fig. 128. Developing ‘‘wing’’ muscles. Transverse section 
through proliferating myoblasts in prothorax, prior to stage 
shown in fig. 130. The column of myoblasts on left of figure is 
growing independently of the larval muscles. The others, which 
are more scattered, are enveloping and drawing together the three 
degenerate larval muscles—eight hours after defaecation (x 800). 

Fig. 129. The same, slightly more anteriorly, showing myo- 
blasts penetrating into the larval muscle, which is shown in trans- 
verse section. A leucocyte is also absorbing part of the muscle 
(x 1200). . 

Fig. 130. One of the two bands of myoblasts in transverse 
section; all larval elements have disappeared—eighteen hours 
after defaecation (x 800). 

Fig. 131. The same, four hours later; the fine syncytial 
columns are beginning to form (1000). 

Fig. 132. Diagrammatic section through the thorax, to show 
the relative size and positions of the two muscle bands (x 40)— 
fresh pupa. é 

Fig. 133. The same, four and a*half days later. Note dis- 
appearance of fat-body, decrease in diameter of oesophagus, and 
increase in size of muscles (x40). 

Fig. 134. A muscle band in transverse section. More cells 
have become merged into the syneytium—four-hour pupa (x 500). 


Pratt XXVI. 

Fig. 185. Longitudinal vertical section through the anterior 
end of the developing ‘“‘wing” muscles—fresh pupa (x500). Note 
three of the syncytial columns (s) applied to the integument, 
whose cells are now dividing to form the insertion cells. 

Fig. 136. The same, eight hours later (500). Note the 
long insertion threads, consisting of two cells generally. At x an 
integumental cell is elongating. 

Fig. 137. The same, sixteen-hour pupa.’ The threads have 
Oe of and are unicellular, and shortening has commenced 

x : . 

Fig. 137a. The same, thirty-six hour pupa. The insertion 
cells have completely retracted (x 500). 

Fig. 138. The same, four and a half day pupa (mature). 
The insertion cells have now broken up into fibrils, on each of 
which a sarcostyle is inserted (x 1200). 

_ Fig. 189. This shows five myoblasts each giving off the fibrils 
which, growing into the syncytial mass, become the sarcostyles. 


524 


The drawing is from a section which just cut the free edge of the 
developing muscles, and the syncytial column has not been repre- 
sented—sixteen-hour pupa (x 1500). 

Fig. 140. Intestine of adult larva (x60). 

Fig. 141. Extremity of lateral hepatic caecum of adult larva 
(x 200). 

Fig. 142. One of the two salivary glands of adult larva 
(x 200). 

Fig. 143. Longitudinal section through the rectum of larva 
in first instar (x 500). Note the blindly ending midgut and rectum. 

Fig. 144. Transverse section through midgut—defaecating 
larva. Two cells undergoing globular degeneration are seen. A 
single replacing cell is visible (x800). 

Fig. 145. The same, four hours later (x1200). Notice the 
shrinking in size of the larval cells and the proliferation of the 
replacing cells. 

Fig. 146. The same, eight hours after defaecation; one side 
of the intestine is shown in section. Note the hepatic caecum, 
and the retention of larval cells in the intestinal epithelium 
adjacent to it (x500). 

Fig. 147. The same, sixteen hours after defaecation. The 
whole intestine is shown in section. Note the drawing in of the 
hepatic caeca through the walls of the intestine as the last remains 
of the dead cells are forced inwards (4500). 

Fig. 148. The same. The hepatic caecum is being sucked in 
(x 500). 

Fig. 149. Section through salivary gland, in larva about to 
pupate (500). Note the globular degeneration. 

Fig. 150. Portion of malpighian tube of imago (x1200). 

Fig. 151. The same, developing in defaecating larva (x 1200). 

Fig. 152. Longitudinal section through junction of oesop- 
hagus and midgut, showing the proliferating cells of the imaginal 
disc of foregut—defaecating larva (500). 

Fig. 153. The same, four hours after pupation. Note leuco- 
cytes attacking the disintegrating temporary midgut (x500). 


Pruate XXVIII. 


Fig. 154. Median longitudinal section through three-hour 
pupa, to show position of the various structures (x80). Note 
the regenerated temporary midgut (m.g.) ending blindly at either 
end, and filled with the débris resulting from the degeneration of 
the larval midgut. The oesophageal cone (0. cn.) is also seen. 

Fig. 155. Longitudinal section along the junction of the 
stomach and developing crop, gizzard, etc., after the front half of 
the midgut has disintegrated—eight-hour pupa (x 4500). 

Fig. 156. The same, thirty-six hour pupa. Note differentia- 
tion of crop, gizzard, and ‘‘drum-shaped chamber’’. ( x 500). 

Fig. 157. The same, a little later, showing the opening of 
the foregut at last into the stomach (x 1000). 

Fig. 158. Small portion of adult oesophagus; note the deli- 
cate spindle-shaped cells (x 1000). 

' Fig. 159. A small portion of wall of crop, to show the folded 
‘“‘napery’’ cells (x 1000). 

Fig. 160. Section through giazzard of adult (x 1000). 

Fig. 161. Section along the rectum of fresh pupa, showing 
developing rectal glands (x 800). 


525 


Vig. 162. Section through anterior end of rectum in larva 
twelve hours before defaecation. Note the malpighian tubes com- 
mencing to grow out (x500). 

Fig. 163. Section through one of the rectal glands—twenty- 
one hour pupa (x500). 

Fig. 164. Section along a rectal gland of imago (x 800). 

Fig. 165. The same; only the thread-like basal cells repre- 
sented (x1500). 

Fig. 166. A nucleus from the rectal gland. 

Fig. 167. Epithelium of small intestine of imago (x 1500). 

Fig. 168. Epithelium of rectum of imago (x1500). 

Fig. 169. Section along a lateral interstinal gland of defae- 
cating larva (x800). 

Fig. 170. Transverse section of the same, sixteen hours 
later, under going degeneration (x 1000). 

Fig g@. 171. Portion of dorsal-abdominal gland from four and 
a half day pupa. In this condition it persists throughout life. 


PuateE XXVIII. 

Fig. 172. The same, in fresh pupa (x 800). 

Fig. 173. Transverse section through the anlage of the same 
on oe postero-dorsal region of the midgut, first larval instar 
(x 1200). 

Fig. 174. Fat cell from thirty-six hour pupa (x800). The 
heavy dots represent the basic storage products; the lighter ones 
the eosinophilous substances. Eight fat globules are seen. 

Fig. 175. Several oenocytes which have been invaginated 
from the ectoderm-larva in first instar (1200). 

ress 1(6, | LT: Disintegrating oenocytes—fresh pupa 
(x 1000). ; 

Fig. 178. Three leucocytes attacking a degenerate oenocyte 
(x 1000)—fresh pupa. 

Fig. 179. Oenocyte in the imago (x 1200). 

Fig. 180. Female reproductive organs—imago. 

Fig. 181. Termination of left accessory gland in longitudinal 
section (cf. fig. 180) (500). 

Fig. 182. Portion of right accessory gland (x 800). 

Fig. 183. One cell from same, to show glandular structure 


Fig. 184. The “lubricating gland”’ of ovipositor in section 
Fig. 185. Section through left half of body of larva in first 


instar in region of testis (x500). 
Fig. 186. Structure of testis in first larval instar. Numerous 


‘spermatogonia and one supporting cell are seen (x 1500). 


Fig. 187. Structure of ovary—defaecating larva. Note some 
incompletely divided oogonia (x 1200). 
Fig. 188. Scent gland of female, opening on body surface 


~ (x 500). 


Fig. 189. Section along an ovarian tubule in four and a half 
day pupa. Note the formation of clusters of nutritive and follicle 
cells. No ovum is as yet recognizable. Most of the distal cells 
are in mitosis (x 1000). 

Fig. 190. Small part of ovarian tubule of imago, showing 


nutritive cells, follicle cells, ovum, and a polar body (x 500). 


Fig. 191. The same (x 350). Note alternate ova and nutri- 


tive cells. 


526 


Fig. 192. Section through end of ovary (fresh pupa). Note 
fourfold connective tissue ingrowth (x350). 

Fig. 193. Two larval leucocytes (x 1500). 

Fig. 194. Leucocyte from eight-hour pupa, showing great 
vacuoles (x 1500). 

Fig. 195. Leucocyte in eight-hour pupa, which has engulfed 
numerous minute pieces of larval tissue (x 1500). 

Fig. 196. Dividing leucocyte, sixteen hours before pupation 


Fig. 197. A leucocyte which has engulfed several large strips 
of larval muscle (2000). 

Fig. 198. Leucocyte that has engulfed larval tissue (x 2000). 

Fig. 199. As in fig. 197 (x2000). 

Fig. 200. ‘‘Normal’’ leucocyte, adult larva (x 2200). 

Figs. 201, 202, 208, Three leucocytes which have absorbed 
pieces of larval muscle and are degenerating among the ‘‘wing”’ 
muscles (x 2000, «1500, x 1500, respectively). 


PruatE XXIX. 


Fig. 204, a, b, c. Three gorged leucocytes which, after feed- 
ing, have ‘‘retired’’ to a cavity of a hollow appendage—eight-hour 
pupa (x 2000). 

Figs. 205, 206. Similar leucocytes, which could not survive 
their meal, and are disintegrating—-eight-hour pupa (1500). 

Fig. 207. A leucocyte in four and a half day pupa degen- 
erating (x1500). 

Fig. 208. A leucocyte of imago, which is recovering from 
its meal—fifty-two hour pupa (x1500). 

Fig. 209. A leucocyte from four and a half day pupa which 
has entirely recovered (x 1500). 

Fig. 210. A nucleus from integumental cell of defaecating 
larva, showing crystals within the great nucleolus (x2300). 

~ Fig. 2010! Heart’ ot adult larva, (420) 

Fig. 212. Posterior end of same, showing ostia (x 800). 

Fig. 218. Section through heart and adjacent structure of 
larva twelve hours before defaecation. Note the delicate cellular 
pericardium outside the heart. Note also the proliferating anlage 
of dorsal abdominal glands (cf. fig. 173) (x 800). 

Fig. 214. Median longitudinal section along dorsal body wall, 
to show metamorphosing ‘‘heart.’? The integument and larval 
cuticle are also shown. The small imaginal and hypertrophied 
larval cells are easily distinguished. Note especially the prolifera- — 
tion of cells along ventral portion of heart. A few leucocytes 
(1) are lying within the heart—defaecating larva (x 1000). ; 

Fig. 215. The same. Numerous embryonic cells extendin 
upwards in a solid column, and overwhelming the hypertrophi 
larval cells (x1000). ~ / i 

Fig. 216. The wall of heart, showing the large larval cells | 
and small embryonic cells advancing upon them (x1000)—defae- | 
cating larva. ) | 

Fig. 217. Section through dorsal body wall of larva six hours 
after defaecation, showing embryonic cells at base of regenerated | 
heart tube (x 500). 

Fig. 218. The same, in a ‘more anterior part of the heart. 
eae on embryonic cells enveloping the regenerated heart tube | 

x ‘ | 

Fig. 219. Regenerated heart (aorta) from fresh pupa (x 1000). 


} 


9 
tio 


527 


Fig. 220. Posterior (contractile) part of dorsal vessel (heart), 
showing the striated muscle cells. 

Fig. 221. <A peripheral nerve from posterior end of defae- 
eating larva. Note larval imaginal elements in the splanchno- 
pleural nerve sheath (x 800). 

Fig. 222. The same in metamorphosis. Note dividing im- 
aginal cell (x 2000)—defaecating larva. 

Fig. 223. The same, showing globular degeneration of the 
fibres within the nerve ( x 2000)—defaecating larva. 

Fig. 224. Nerve cell from brain of imago (x 1500). 

Fig. 225, a, b, c, d, e. Stages in the fusion of ganglia in’ 
the ventral nerve cord during the early hours of pupation. Semi- 
diagrammatic. 

Fig. 226. A regenerated nerve ending on a muscle fibre in 
the rectum—four and a half day pupa (x 1000). 


PLraTE XXX. 

Fig. 227. Longitudinal section through a metamorphosing 
thoracic ganglion, from defaecating larva. Note masses of dead 
tissue. Several imaginal cells are dividing by mitosis (x 900). 

Fig. 228. Five nerve cells from thoracic ganglion, eight 


hours after defaecation. Two are developing nerve fibres 


(x 1500). 

Fig. 229. Metamorphosing neuroglia network of ventral nerve 
cord (x900). 

Fig. 230. Transverse section through brain of first larval 
instar, showing larval and imaginal elements (x700). The 
apparently greater size of the imaginal cells is due to the fact 
that most of the cell substance in the larval cell is contained in 
the nerve fibres. 

Fig. 231. Dorsal view of brain of adult larva (x190). 

Fig. 232. The same, from fresh pupa. Note development of 
optic ganglia, and the migration of first ganglia of ventral nerve 
cord (x190). : 

Fig. 233. Brain of pupa, twenty-six hours old, view from in 
front and from slightly below. Note incorporation of first ventral 
ganglion into the brain (x190). 

Fig. 234. Brain of imago, view from front. The right hemi- 
sphere is represented as if the anterior portion were dissected. 
away, revealing the nerve strands at the rear of the brain (190). 

Fig. 235. Section through metamorphosing brain tissue 
(eight hours after defaecation (cf. fig. 77). Notice the mass of 
dead tissue; also the degenerate nerve strand. Imaginal elements 
are proliferating (x1160). 

Fig. 236. Transverse section through half of the brain 
{twenty-four hour pupa), showing regenerated nerve strands, 
ganglia, etc. (x190). 

Fig. 237. Neuroglia network within ocellar nerve strand of 
brain—twenty-four hour pupa (x900) (cf. fig. 236). 


528 | : | 


A PRELIMINARY NOTE ON THE FOSSIL WOODS FROM 
SOME AUSTRALIAN BROWN COAL DEPOSITS. 


By E. Dororuy Nosss, B.Sc., 
Research Scholar in Botany, University of Adelaide. 


[Read October 19, 1922.] 


The South Australian brown coal deposits at Moorlands 
have recently been the subject of geological investiga- 
tion,“) but the plant remains they contain have not as yet 
been examined botanically. It therefore seemed that a 
study of the plant constituents of this material might prove 
of interest. A quantity of material was made-available for 
investigation by Mr. A. C. Broughton, manager of the Moor- 
lands Mine, whom I take this opportunity of thanking. 

For comparison with the material from Moorlands a col- 
lection of woody fragments was obtained from the brown coal 
deposit uncovered this year at Yallourn, Gippsland, Victoria. 

My thanks are due to Professor Osborn, Department of 
Botany, University of Adelaide, for the interest he has shown 
throughout the course of the work. It had been intended to 
carry the investigation further, but, circumstances having 
arisen that will prevent my doing so, the results are given 
here as far as they go. 


MATERIAL FROM MOORLANDS. 


The coal consists of a matrix in which are embedded frag- 
ments of lignite often of considerable size. Occasional layers 
of black shiny coal are noted, also lumps of a jet-like sub- 
stance thought to be resin. Several specimens of coal show a 
laminated structure, between the layers of which are numerous 
leaf cuticles and other leaf fragments, all of which appear 
to be Dicotyledonous types. It was, however, thought advisable 
to confine the present investigation to an examination of the 
structure of the ‘‘wood,’’ since it has been shown that leaf 
impressions and leaf cuticles alone are of doubtful value) in 
generic determinations of Tertiary plants. The woody 
material was separated out from the matrix, and after suit- 
able treatment it was found possible to obtain good microtome 
as well as free-hand sections. 

Forty-five distinct pieces of wood were obtained. These 
show varied states of preservation, but a large number were 


(1) Broughton, A. C.; also Mawson, Sir D., and F. Chapman. 
(2) Broughton, A. C., l.c., p. 252. 
(3) Seward, vol. iv. Andrews, 1916. 


529 


sufficiently well preserved to be determinable. Of the 45 
fragments, 23 showed Coniferous characters, while 4 proved 


to be Angiosperms. 


The remaining 18 fragments were too 


poorly preserved to allow sectioning. The 4 Angiosperms 


3 
Sf % 


Mesembrioxylon, sp. 
Moorlands A. Radial 
section showing the 
tracheal pitting. 
x 275, 


aT) 


vvvvagell ui 


spa peyet TON 


oyKTo) 


Vey ee i 

~~ 

O et Fey: 

~ wey Be spe 

cS ee pe / 
' 


es 
ta 
A\Vn 1 


sete treny 


a 
ppb beas etayh 
7 
sa ALLEL 
if 


U2 vy yt 
SUN OTT Tn 


Fig. 3. 
Mesembriozylon, sp. Moorlands 5. 
Radial section showing trachea! 
pitting. x275. 


lands 
showing 


Fig. 2. 
Mesembrioxylon, sp. Moor- 


Radial section 


the pits in the 
field. x275. 


have not given sufficient data to admit of identification, 


though it is probable that there are two species represented. 
_ Of the 23 fragments definitely Gymnosperms, only two have 
structural! features sufficiently preserved for detailed de- 
scription. 


530 


DESCRIPTION OF SPECIMENS. 
MESEMBRIOXYLON, sp. 
Moorlands A (figs. 1, 2). 
The state of preservation is good. The lumen of the 
tracheids is not obscured by brownish matter nor do the rays 


contain any dark-brown substance. 
Growth rings are clear. 


Bordered pits uniseriate, separate, and circular (fig. 1). 
Medullary rays aniseriate, two or three cells high, occa- — 
sionally four or five. The cells are ae from dark- 


= aa “OD [| OD [I] @2 | DO 
2b am 9 if sallKc 
amg (281 (tata 


= Pu. | sain 


S 


Fig. 4. 
Mesembrioxyion, sp. Yallourn A. Radial 


section showing pits in the field and 
tracheal pitting. x 275. 


brown contents. There may be one or two oval, oblique, 
apparently simple pits in the field (fig. 2). The presence of 
pits on the horizontal and tangential walls of the rays is well 
known to be a difficult character ey determine, (4) and negative 


(4) Seward, call iv., p. 169. 


‘< 
:. 
} , 


531 


evidence is unsatisfactory; however, pits do not seem to be 
present. 


No rims of Sanio are apparent. 


MESEMBRIOXYLON, sp. 
Moorlands B (fig. 3). 
A considerable amount of compression has taken place in 


certain regions of this specimen. The medullary rays and the 
lumen of the tracheids are filled with brown contents. 


Mesembrioxylon, sp. Yallourn B. 
Radial section showing pits in the 
field and pitting on the radial and 
oblique walls ey the tracheids. 
x 275. 


| 

| 

| 4 

a Fig. 5. 


Growth rings are present, 

Bordered pits uniseriate, separate, and circular (fig. 3), 
only showing in regions of better preservation. 

Medullary rays uniseriate, two or three cells high, rarely 
four or five. Many ray cells have dark-brown contents. 


Le '; | 
y 1 
\ { 
4 


The pitting in connection with the medullary rays is 
not preserved. 
Rims of Sano not observed. 


Se 


532 


It would appear that there is a close affinity between 
the two specimens. A resemblance to the living genus 
Callitris is evident, but until further material and more 
data are available it seems inadvisable to definitely imply an _ 
affinity with a particular existing genus purely on the evid- 
ence of the wood. it seems, from the characters described 
for both specimens, that they are best included under the 


genus Mesembrioxylon instituted by Professor Seward.) 


Fig. 6. 
Mesembrioxylon, sp. Yal- 
lourn B. Tangential sec- 
tion showing medullary 

rays. Xoo. 


MATERIAL FROM YALLOURN, GIPPSLAND, VICTORIA. 


This material is in a much better state of preservation and 
exhibits several features of interest. 

Kleven different wood specimens from Yallourn have been 
examined, but only sufficiently complete data have been 
obtained to justify description of four. One Angiosperm 
was recognizable, while the remainder were undoubtedly 
Gymnosperms. 


(5)Seward, vol. iv., p. 208. 


533 


DESCRIPTION OF SPECIMENS. 
MESEMBRIOXYLON, sp. 


Yallourn A. (fig. 4). 

This wood shows well-defined growth rings. Tracheids 
or xylem parenchyma containing dark-brown matter were not 
observed. 

Average diameter of the tracheids is 45y. 

Bordered pits are in one or two rows, contiguous but not 
compressed. Where the pits are in two rows they are alternate 
or subopposite (fig. 4). Pits occur occasionally on the 
tangential walls of the tracheids. 


atte 


~ 


Veeperpae 


Stage 


afm au 


Bee.) 7 
Cupressinozylon, sp. Yallourn. Tangential 
section showing medullary rays and _ occa- 
sional pitting on tangential walls of tracheids. 


> 


xia. 


ee 


Medullary rays simple, uniseriate, usually from one to 
eight cells high, occasionally ten. Some of the ray cells con- 
tain dark-brown matter. There are from one to four circular 


534 


pits im the field in connection with the rays. These pits have 
narrow oblique slits (fig. 4). . 
Spiral bands appear on the tracheids, but no rims of 
Sanio were observed. 
Apparently there is no zylem parenchyma. 


MESEMBRIOXYLON, sp. 
Yallourn B (figs. 5, 6). 
Growth rings are well defined in this wood. No tracheids 
or xylem parenchyma with dark-brown contents were found. 


Fig. 8. 
Cupressinoxylon, sp. ; 
Yallourn. Tangential Fig. 9. 
section showing Cupressinoxylon, sp. Yallourn. 
medullary rays. Radial section showing pits in the 
x 50. field and pitting on the tracheids. 


x W75. 


The average diameter of the tracheids is 30uy. 

Bordered mts are circular, separate, and scattered, 
usually in one row. Occasionally the pits are slightly 
flattened (fig. 5). 

Medullary rays are uniseriate, from one to eight cells 
high, occasionally more than eight cells (fig. 6). The ray 
cells are large and thick-walled. The pitting in con- 
nection with the medullary rays is well preserved, there being 
one large oblique pit in the field (fig. 5). Occasionally the 
pit may have the appearance of a border. : 


535 


CUPRESSINOXYLON, sp. 
Figs. 7-9. 

Growth rings are well defined here. Many cells have the 
lumen filled with dark-brown matter. The medullary rays 
are close together and have light-brownish contents with 
occasional ‘“‘resin-spools.’’ 

The average diameter of the tracheids is 20n. 

Bordered pits are uniseriate, separate, and circular. The 
spring wood may show two rows of pits on a few tracheids. 
Pits also occur scattered on the tangential walls (fig. 7). 

Medullary rays simple, uniseriate, and numerous through- 
out the wood 


Fig. 108° pas 
Dadozylon, sp. Yallourn. Fig. 11. 
Hexagonal pitting on Dadozylon, sp. Yallourn. 
radial walls of the tra- Numerous small pits in ' 
cheids. x 275. the field. x 275. 


cells high. At times they may reach a height of thirty cells. 
Some of the ray cells have contents. The pits in the field 
are one or two oval and oblique, and there may be the appear- 
ance of a border (fig. 9). 

Aylem parenchyma is present with resinous(?) contents. 

The general characters, especially the pitting, in con- 
nection with the medullary rays, and the presence of wood 
parenchyma, suggest that this specimen should be included 
under Cupressinoxylon. 


DaDOXYLON, sp. 
Figs. 10, 11. 
This is a roughly cylindrical piece of wood, oval in cross 


_ section, with the outer cortex preserved and remaining 
_ attached to the woody cylinder. The long diameter of the 


536 ji 


whole is about 2°5 cm... The cell detail of the specimen is not 
well preserved, but the following details can be determined : — 

In transverse section many cells have the lumen filled 
with brown matter. Occasional irregular strands of paren- 
chyma occur in the wood in places, but their nature could not 
be determined. 


Growth rings are present. 

Bordered pits on the radial walls of the tracheids are 
in two or more rows, alternate, compressed, and hexagonal. 

Medullary rays simple, uniseriate, from one to eight cells 
high. Radial sections show numerous small oval pits in the 
field. Horizontal elements filled with dense dark-brown sub- 
stance cccur in association with the rays. : 


CoNncLUSION. 


It would appear that considerable Gymnosperm forests 
have contributed to the formation of the brown coal deposits 
of Moorlands, South Australia, and Yallourn, Victoria. Such 
forests do not occur in South Australia or Victoria to-day 
though there are occasional open forests of Calhtris growing 
under semi-arid conditions. No mixed Coniferous forests 
exist -in these parts, yet these must have given rise 
to the Yallourn lignites which have yielded several distinct 
types of Gymnosperms, namely, two species of dJ/esem- 
brioxylon, one Cupressinoxylon, and one Dadoxylon( ? ). 
The Moorlands forests would appear to have been more uni- 
form, so far as preservation of the wood allows one to say. In 
view of the abundance of Dicotyledonous leaf fragments from 
Moorlands, the scarcity of Dicotyledonous wood is strange. 


REFERENCES. 

Brovucuton, A. C.—‘‘Notes on the Geology of the Moorlands 
(S.A.) Brown Coal Deposits,’ Trans. Roy. Soc. S. Austr., 
xlv., pp. 248-253, 1921. i 

Mawson, Sir D., and F. Cuapman—‘The Tertiary Brown 7 
Coal-bearing Beds of Moorlands,’’ ibzd., xlvi., pp. 
131-147, 1922. 

SeEwarD, A. C.—‘‘Fossil Plants,’ vol. iv., Cambridge, 1919. 

AnpDREws, E. C.—‘‘The Geological History of the Australian 


Flowering Plants,’ Am. Journ. Science, vol. xlii., pp. 
171-232, 1916. 


537. 


ON A NEW GENUS AND SPECIES OF 
AUSTRALIAN LYCAENINAE. 


By Norman B. TInDALE. 


(Contribution from the South Australian Museum. ) 
[Read October 19, 1922.| 


Pyare XXXL. 


ADALUMA, n. gen. 


Forewing with vein 11 parallel with vein 12, vein 6 arising 
with vein 7, veins 1 to 7 (in male) bordered discally with 
black scales; costa strongly arched; termen well rounded. 
Hindwing evenly rounded, apex of cell acute, vein 3 arising 
some distance below apex of cell. Beneath white, with 
terminal series of dots. Antennae short, less than half ex- 
panse of wings. Hyes smooth, palpilong. Type, 4. wrumelia, 
from the Northern Territory. 

Allied to Candalides, Hubner, to which it is similar in 
venation. The shape of the cell of hindwing differs from C. 
zanthospilos, Hubner, in being more acute at apex. The 
antennae are extremely short; in this character it resembles 
the peculiar genus Vesolycaena, from which otherwise it is 
distinct. 

The names chosen, ‘‘adaluma’’ and ‘‘urumelia,’’ are two 
native (Nungubuyu tribe) words meaning ‘“‘flowing stream’’ 
and “‘butterfly.”’ The butterfly was first taken on the banks 
of the Roper River by a native, at the aborigines’ reserve, 
which is over 70 miles from the sea. 


ADALUMA URUMELIA, 01. sp. 

d. Above. Forewing silky-white tinged with blue; 
apex and termen narrowly grey-black, veins tipped black at 
termen ; veins 1 to 7 bordered with black scales in discal area. 
Cilia black, tipped with white. Hindwings silky-white, a 


terminal line black. Cilia black, tipped with white. 


Beneath. Silky-white. Forewing with two large terminal 
black spots in areas la and 2; terminal line black. Hind- 
wing with a terminal series of round black dots, terminal 
line black. Cilia black, tipped with white, at dorsum white. 
Antennae short, well clubbed, joints short, black, tipped with 
white; palpi long, tipped with black. Expanse, 30 mm. 


538 


Loc.—Northern Territory: Roper River, March, 1922 
(Mrs. H. E. Warren and a native). Type, I. 13771. 

This species is known from two males, one fragmentary, 
but the other perfect. Following Waterhouse and Lyell,® 
in a hnear arrangement, it would be best placed between 
Nesolycaena and Philiris. 


EXPLANATION OF PLATE XXXI. 


Fig. 1. Adaluma urumelia, male, upper-surface, x3. 
Renee ie i .5  under-surface, x3. 


(1) Waterhouse, G. A., and Lyell, Butterflies of Australia, 
p. 76, 1914. 


539 


ON THE ECOLOGY OF THE OOLDEA DISTRICT. 


By R.S. Apvamson, M.A., B.Sc., and T. G. B. Osporn, D.Sc. 
[Read October 19, 1922.] 
PuatTeS XXXII. ro XXXVI. 


Comparatively little has been written about the ecology 
of the arid regions of Australia, though these form a large 
portion of the continent. In South Australia, which has an 
area roughly three times the size of the British Isles, five- 
sixths of the total area has under 10 in. of rain per annum. 
A vast field, therefore, is awaiting examination. One reason 
that has contributed to the neglect of this work is the diffi- 
culty of visiting the places and the time occupied in the 
journey. The recent opening of the Transcontinental (Hast- 
West) Railway connecting South and Western Australia has 
made very accessible an area that until the last few years was. 
visited by but few white people. The Ooldea district lies well 
within this area, and offers scope for examining the arid flora, 
the more so that, owing to an abrupt change in the type of 
soil in the immediate locality two distinct habitats are avail- 
able. Cannon visited Ooldea recently in connection with his 
work on the Arid Flora of South Australia, but his account 
was brief, and no attempt was made by him to deal with the 
flora as a whole. 


Ooldea is a station on the Transcontinental (Hast-West) 
Railway, 427 miles west of Port Augusta and 374 ft. above 
sea level. The Ooldea district is one of great biological 
interest because of its situation on the eastern boundary of the 
Nullarbor Plain, at the point at which the railway line leaves 
the sandhills and runs over the limestone plains. In August 
of this year we made a stay of six days in the district, and as 
a result of this visit the following account of the vegetation 


is given. It is a pleasure to express our thanks to the Pre- 


sident and members of the Ooldea Progress Association for 
the facilities they placed at our disposal; to Mr. T. Davison, 
engineer at the “‘Soak,’’ for his guidance in that area; and 
to Mr. A. G. Bolam, stationmaster at Ooldea, for meteor- 
ological data and help in various ways. We are also indebted 
to Mr. J. M. Black, who has determined some of our material 
for us. Mr. J. H. Maiden, F.R.S., has kindly named the 
Eucalypts. 
Q2 


540 


PHYSIOGRAPHIC. 


The Nullarbor Plain is an area of limestone country 
extending westward from Fowler Bay to beyond Eucla in 
Western Australia. Its east-west extent is 450 miles and 
its greatest north-south width about 200 miles. The area 
within South Australia is 17,767 square miles, throughout 
the whole of which there is no watercourse or lake. It is 
underlain, however, by the Eucla basin of artesian water 
which has been tapped by bores at depths varying from 
298 ft..to 1,500 ft. A bore sunk at Ooldea gave water at 
480 ft., containing 7°716 ozs. of salt to the gallon.) 

Little has been written on the geology of the area, the 
account by Tate,@) who visited the seaward margin of the 
Nullarbor Plain in 1879, being still the most complete. 


~The Nullarbor Plain forms a part of the Bunda 
Plateaw, which, according to Tate, is the ‘‘elevated bed of 
the older Tertiary sea, the sediments of which were deposited 
within a very extensive granitic basin.”’ These igneous rocks 
come to the surface at various places round the edge of the 
basin, an outcrop occurring to the east of Ooldea. According 
to Howchin ©) the limestones are Miocene (Janjukian), over- 
lain by older Pliocene (Kalimnan), at Wilson Bluff, near 
Eucla. 

The surface of the plain is not perfectly level, but rises 
and falls in gradual undulations.(*) The limestone is covered 
by a red sandy loam varying from a few inches to a foot or 
more in depth, which soil, being fine grained, bakes hard 
when dry. Fragments of limestone, however, are freely 
interspersed with the soil, and appear at the surface in most 
places, especially on the ridges. 

Here and there are ‘‘dongas,’’ or slight depressions vary- 
ing in area from less than an acre to some hundreds of acres. 
The dongas are said to be the largest at the western side of 
the plain. Those seen by us were small; they are said to be 
infrequent in the centre of the plain. The soil in the dongas 


> 


(1) Rep. 3rd Interstate Conference on Artesian Water, Ade- 
laide, 1921, p. 16. Adelaide, Govt. Printer, 1922. 

(2) Tate, R., The Natural History of the Country around the 
Head of the Great Australian Bight, Trans. and Proc. Philos. 
Soc. of Adelaide, S. Austr., i1., pp. 94-128, 1879. 

(3) Howchin, W., ‘‘Geology of South Australia,’ pp. 457 and 
466, Adelaide, 1918. 

(4) A quantity of interesting information as to the Nullarbor 
Plain is collected as a supplement to the Presidential Address to 
the South Australian Branch of the Royal Geographical Society, 
en 1917-18. Proc. Roy. Geol. Soc. S. Austr., xix., pp. 101-158, 


541. 


is sandy and of greater depth than on the plain, and, in those 


seen by us, free from limestone. In addition to the dongas 


there are numerous “‘blow holes,” fissures of varying depth 
in the limestone, that open in many cases into caves below. 
It seems probable that the dongas represené areas of sub- 
sidence.©) The whole structure of the plain is such that the 
rain which falls readily disappears below the surface. Only in 


_ the dongas is there sufficient soil to hold an appreciable water 


reserve; the soil of the plains is too shailow. 
Starting at Ooldea and running eastwards for about 50 


miles is a sandhill region, consisting of a series of ridges of 
-red-coloured sand with flats between. The ridges, which may 


be as much as 30 ft. in height, run approximately north-west 
by south-east ‘at Ooldea, but there are many irregularities 


and connecting ridges. The sand forming the ridges is rela- 


tively stable and no drifting of large masses occurs. Indeed, 
some, at any rate, of the larger ridges have a core of travertine 
limestone 3 ft. or so below the surface. The soil of the inter- 
vening flats is generally level and composed of finer particles, 
and is consequently firmer. This soil, on the whole, appears 


deeper than on the ridges themselves. Whatever be the 


source of the sand, it is clear that a certain amount of wind 
sorting has taken place, with the result that over the whole 


area two habitats differing in their edaphic conditions have 


been developed. 


About three miles north of Ooldea lies the famous Ooldea 
Soak. This is a shallow basin, 10 acres or more in extent. 
The sand of the Soak and.its surrounding ridges is whiter 
and less stable than that of the majority of the sandhills; 
it rests upon an underlying layer of bluish clay. Within the 
drea there is considerable drift, so that the floor of the basin 
is-broken by various hollows and ridges held by shrubs. Below 
the surface, at a depth of 5 ft. to 15 ft. or more, lies a water 
table, the water being fresh, somewhat alkaline, but generally 
quite potable. The Soak has been the scene of human activi- 
ties for a great period of time, for it was known to the 
aborigines long before the expiorer, E. Giles (who was one of 


the first whites to visit it), used it as a base camp in 1875. 


Later it was used as a centre for sheep grazing,“ but appears 


to have been abandoned for that purpose some time before 
the building of the Transcontinental Line made it a place of 


6) Brown, H. Y. L., quoted Proc. Roy. Geog. Soc. S. Austr., 


Hoc. cit., p. 134. 


(6) Giles, E., Australia twice Traversed, ii., p. 152, 1889. 
(7)Brown, T., Proc. Roy. Geog. Soc. S. Austr., loc. cit., p. 149. 


- 


542 


importance. At present over twenty wells have been put 

down within the area, and water is pumped to Ooldea at the 

rate of 10,000 gallons a day. it is legitimate to assume that 

the vegetation within the Soak basin has suffered somewhat 

from human interference, and it may suffer still more if 

pumping permanently lowers the water level. Mr. Brown, 

referring to his visit about 1887, speaks of ‘‘clumps of bull- 

- rushes’’ growing here and there. None are there now, while 

of the plants collected by us only Adriana tomentosa—locally 
called “‘water-bush’’—seems definitely tndicative of consider- 

able edaphic water. Within half a mile of the Soak, in a 
north-easterly direction, lies a salt lake or claypan, smaller 

in area than the Soak itself. This was dry at the time of 

our visit, but the friable soii overlying the claybed was 

impregnated with crystals of soluble salts, including gypsum, 

that caused a white efflorescence at the margins. 


CLIMATE. 


Climatological data for Ooldea were not kept before 
December, 1916, because, until the construction of the Hast- 
West Railway, there was no settlement at the place. From 
that date rainfall records have been kept, the temperatures 
have been recorded less regularly. The actual figures cover 
too short a period to allow of generalization on them ‘alone, 
but, taken in conjunction with what is known of other places 
similarly situated, they are useful. 


The following table gives the rainfall in inches from 
January, 1917, to October, 1922: 


1917. 1918. 1919. 1920. 1921. 192m) 


January Big | 0°00 0-70 0-00 0-03 0:00 
February nh AS 0°00 1-63 0-00 2°83 0°59 
March Boe ist 1°00 0-00 0-08 0:28 0-71 
April Reha gy! 0°60 2-11 1-00 0-01 1-279 
May rie One 0°63 0-06 1-39 2°13 0°25 
June sie bag 1:06 0-25 0-45 0°91 O74 — 

July APLAR 0°16 0-16 0-68 0°15 0°25: 
August ihe PBZ 139 0-57 0-69 0-15 0:00: 
September... 0°76 0°29 0°08 2°32 0-03 0-00. 
October ere GS) 0°72 0:54 0-64 0-36 0-15 
November et bee |G hes 1 0-20 0-49 0°93 — 
December gg Nees 0°39 0°33 0-42 0°35 —_ 
Total for Year 14°11 pale 6°63 8-16 8-16 *3°96: 
Wet Davyve.teoe 65 30 27 AT 43 *2F 


*Totals for 1922 on figures for 10 months only. 


bay 


as, 


543 


The monthly average temperatures in degrees F. at 
9 a.m. and 4 p.m. are given below, no maximum and minimum 
readings being available :— 
Jan. Feb. Mar. April May June July Aug. Sept. Oct. Nov. Dec. 
Sam. 70 6 7 5d 52 45 48 49 57 58 6 75 
mm, 105 6 104:«CO99 «= 690) 89 5 SCD 8Ss«*2108.s «106 
The highest shade temperature recorded is 125° F., which 
occurred in January. 
The rainfall is not only low but erratic. Though, 
broadly speaking, the summer is the dry season and the winter 


the wet one, the falls at best are too low to make the winter 


always a season of active vegetation, whilst a heavy downpour 


‘In the summer may have a marked effect in producing con- 


siderable activity on the part of some of the annuals at least. 
Owing to the very porous soil the rain sinks into the ground 
as it falls, there being no sign of a watercourse in the district. 

Another feature in regard to the rainfall is the amount 
that falls at any one time. Though the monthly total may 
seem relatively high, if it be composed of a number of small 
Showers, as is often the case in arid Australia, the general 
effect on the vegetation is slight. Cannon has drawn atten- 
tion to what he terms effective and ineffective rainfalls in 
arid South Australia, a fall of 0°15 in. in twenty-four hours 
being insufficient to produce more than a surface wetting of 
the soil. This important feature, which is recognized by Aus- 
tralian pastoralists, must be remembered when considering 
the monthly falls at Ooldea or any other station in this part 
of arid Australia. 

The temperature records show the climate to be one of 
considerable extremes. Short periods of great heat, over 
100° F. in the shade, may be expected during six months of 
the year, while the average daily temperature at 4 p.m. 
Nov.-Feb. inclusive is 103° F. or over. The area, however, 
is so open that heat is rapidly lost by radiation at night, 
even in summer, while in winter the diurnal range of tem- 
perature on the soil between-sun heat of the day and at 
night is still more severe. Ground frosts are not uncommon 
on clear nights during three or four months of the year. No 
records of temperatures at the surface of the soil have been 
taken, but it is certain that they must reach very high figures, 
especially during the summer. In August, at the time of 
our visit, on a day on which there was a frost upon the 


_ ground before sunrise, the sand at midday was so hot in the 
sun that it was uncomfortable to touch it with the hand. In 


summer the heat at the surface of the soil must be so great 
as to seriously affect plants unless they are well insulated at 


the ‘‘collar’’ by cork or some other nonconductor. In this 


544 


connection it may be noted that perennial herbaceous plants — 
are noticeably rare in the Ooldea flora. Another adverse factor 
is the drying winds which sweep over the plain. Ooldea is 
80 miles from the sea, and the intervening country is similar 
waterless plain or sandhill. Even the south and south-west — 
winds have their humidity reduced before reaching Ooldea, 
while any other wind travels considerably further overland — 
before it gets to that place. All winds, but especially thoses 
off.the deserts that lie north and north-east and north-west, 
must be regarded as influences supp e adversely to the | 5 
vegetation. 


PREVIOUS WORK ON THE FLORA. 


The paper of Tate before mentioned includes a list of the 
flora at the head of the Bight as well as some notes on the 
vegetation. The region traversed by him, being near the 
coast, has a higher rainfall than Ooldea, and also gains con-— 
siderable moisture from sea mists for a-.distance inland of 
20 miules.(8) Nevertheless, the vegetation seems essentially 
similar. Tate’s(?) notes on the flora are valuable, especially 
noteworthy being the clear distinction that he makes between 
such halophytes as Arthrocnemum spp., and xerophytes as 
Kochia sedifolia. Saltbushes, Atriplex spp., are assigned an 
intermediate position ; they show considerable salt toleration, 
but not to the same degree as the truly halopes shrubby 
Salicornias. 


The Ooldea region has been studied floristically recently 
by Black.0® Cannon visited the district in 1918, and his 
observations are included in his recent work on the “‘Arid 
Portions of South Australia.”’ No attempt is made“) by — 
Cannon to give a complete account of the flora, but the main — 
habitats are studied with reference to some of the most pro- — 
minent plants occurring there. Cannon distinguishes the — 
Nullarbor Plains, the sandhills, and a transition region 
between the two; the dongas and the hollows between the | 
ridges in the sandhills are also indicated as probably dis- | 
tinct. With his account we are in general agreement, except 
that the characteristic bluebushes and saltbushes of the plains — 
are classed by him as halophytes. With this we disagree. 


. | 
(8) Brown, T5 toc. cit., p. 147. 
0) Tate. Ri Mec mit.) pp. 11S-195. 
(10) Black, J. M., Trans. Roy. Soc. S. Austr., xli., pp. 378-3890; 
1917, and xlv., pp. 5-24, 1921. 


(2) Cannon, W. A., Plant Habits and Habitats in the re 
Portions of South Australia, Carnegie Inst. of Washington, Publ. 
No. 308, 1921, pp. 81-89. . 


—) 
= 
; ~- a %s 


545 


VEGETATION. 


At least two very markedly different kinds of habitat 
occur near Ooldea, namely, the Nullarbor Plain and the sand- 
hills. These will be described separately. 


Nullarbor Plain.—The plain itself bears a very open 
vegetation of a highly xerophytic character. The soil being of 
a@ porous nature, and one which does not easily form a dust 
mulch, the water supply for vegetation is scanty and very 
uncertain in amount. This is clearly expressed in the sparse 


_ plant covering. On the plain itself trees or woody plants of 
any size appear to be absent. 


The chief plants are the “‘bluebush,’’ Aochia sedifolia, 
and in rather less quantity the ‘‘saltbush,’’ Atriplex vesi- 
eartum, which occur in communities which are often sharply 
delimited from one another. Other species of Kochia and of 
Atriplex also occur, but in less quantity, ¢.g., A. triptera, 
v. ertoclada, kh. pyramidata, etc. 


These plants, which we may term the “‘character plants,” 
are small bushes, 1 ft. to 2 ft. in height, which stand in most 
parts at considerable distances from one another—often as 
much as two, to three yards (pl. xxxu., fig. 1). In spite of 
this condition, however, in which the plants develop quite 
independently of their neighbours, the communities are 
usually quite pure as regards their character species. Aochia 
sedifolia is much more common than Atriplex, and covers a 
much greater area of the plain. It occupies nearly all the 
surface where the fine-grained red soil occurs, while Atriplex 


vesicartum occurs in those parts where some sand is present or 


the soil is deeper and looser. The latter is especially abundant 
near the eastern margin of the plain and along the line of 


junction with the sandhills. So far as our observations 


extend, Aochia would appear to be more xerophytic, br, at 
any rate, able to withstand drier conditions. This agrees 
with the observations of Tate,“2) who describes the plateau 
near Eucla as covered with rhea while 4 triplex occurs in 
slight depressions around, but not extending into, saline 
swamps. These Chenopodiaceous bushes, which have their 
leaves more or less densely covered with hairs or scales, appear 
white or silver-grey in colour, and, growing as they do in a 
soil that also appears pale owing to the numbers of limestone 


_ fragments, give a landscape at once characteristic and peculiar. 
_ The observer has a sense of uniformity and vastness which is 


unbroken by any marked change in the surface or by any 


sign of life, either bird or animal. 


(12) Tate, eae. Phil. ae aa ee i1., p. 120, 1878. 


546 


The plants have rather small leaves, which are spread- 
ing, and, especially in Aochia sedifolia, distinctly succulent. 
It seems ‘possible that these plants not only store up water in 
their leaves when rain falls, but also possess the power of 
absorbing moisture directly by their leaves by means of the 
hairs or scales. All the residents, both here and in other arid 
parts of the State, remark on the obvious freshening up of 
the bluebush or saitbush that occurs immediately after rain. 


On the other hand, the hairy covering may well be correlated 
with the intense light and heating that the plants in such a 


habitat have to withstand. 

The severity of the conditions for perennial plants here 
is expressed. both in the open nature of the communities and 
also in a striking way by the amount and number of dead 
plants that occur (pl. xxxu., figs. 1, 2). Whole stretches may 
be seen, extending a mile or more, in which all the perennials 
are dead, presumably killed by drought. No evidence of fire 
was noticed, not even in the neighbourhood of the railway 
track, and, except at the margins a the plain, little effect has 
been produced by the ubiquitous rabbit. 

Besides these perennial plants a very considerable number 
of annual species occur on the plain between the bushes of 
Kochi and Atripier. The amount and nature of these 
annuals vary according to the season at which the rain falls, 
and at certain times may temporarily alter the whole general 
appearance. At the time of our visit very little rain had fallen 
for some time previously, and this therophyte flora was poorly 
developed. One of the most abundant ‘annuals was Salsola 
kali, var. strobilifera, which at this season was dead. The 
plant occurs in great abundance in parts, and appears especi- 
ally to spread where the Kochia has been killed off. Other 


very generally distributed annuals were Cephalipterum 


Drummond, Zugophylium ovatum, Angianthus tomentosus, — 


Gnephosis cyathopappa, Goodena pinnatifida, Goodenia 
pusilliflora, Helipteruwm pygmaeum, Lemdium phlebopetalum, 


Echinospermum concavum, Isoétopsis granunfolia, and — 


Tetragonia expansa, also Bassia scleroloenioides, perennial. Of 
these the first two were the most abundant or most prominent. 
Cephalipterum especially, with its white flower-heads, sam 
quite a character to the plains. ; 

At the time of our visit grasses were remarkable for their 


scarcity ; a few scattered plants of Danthonia penicillata and — 


Stipa setacea and Stipa scabra were noticed, but so few that | 


nowhere on the plain was grass vegetation at all prominent. 


At other times these grasses, and especially Stipa, sp., may — 


become a marked feature after good rains. At the time of 
our visit most of the Stipa plants noted were dead. 


547 


Dongas.—These are shallow depressions of varying size 
and extent which occur scattered over the plain. Except in 
the smallest and most shallow, there is a considerably greater 
quantity of soil in them—a soil, too, which is much freer 
from limestone fragments. 
The dongas examined, which, however, were all com- 
paratively close to the eastern border of the plain, bear a 
vegetation with trees or shrubs (pl. xxxii., fig. 1). The most 
general were species of Acacia, especially A. aneura (mulga), 
A. Oswaldu, A. tetragonophylla, with less often A. 
Randelliana. 
Other trees occurring were Pittosporum phillyraeoides, 

Fusanus persicarius, and locally Casuarina lemdophloia. This 
last was seen only in the largest depressions where there was 
most soil. Hremophila Latrobe: occurred rarely. Beneath 
these trees the soil was quite bare in the smaller depressions, 
but in the larger ones herbaceous plants (mostly annuals) 
were present in some quantity. These were generally quite 
different from those on the general surface of the plain; 
Cephalipterum Drummondu, however, was usually present. 
_ Stipa scabra was locally quite abundant, though at this season 
it appeared dead. 3 

Other annuals found in these dongas were Salsola kali, 
Lavatera plebeja, Convolvulus erubescens, Lotus australis, 
var. pubescens, Helipterum. floribundum, Nicotiana suaveo- 
dens, Calandrima volubilis, Pimelea simplex. 

In several of the dongas near the eastern margin of the 
plain much of the undergrowth has been destroyed by rabbits, 
which have their burrows there. 


At the eastern margin of the Nullarbor Plain a certain 
amount of change in vegetation is seen. More soil is pre- 
sent and a richer vegetation develops. Small trees and shrubs 
| occur scattered over the surface, though often localized in a 
peculiar way (pl. xxxil., fig. 2). Most of the shrubs found in the 
dongas grow in this situation, with the exception of Casuarina 
depdophloa. The most abundant shrubs are Acacia aneura, 
A. tetragonophylla, and Eremophila Latrobei, with Acacia 
Randelliana, A. Oswaldui, Pittosporum  phillyraeoides, 
FPusanus persicarius, and Eremophila oppositifolia, less fre- 
quently. Here also the undergrowth is rather more luxuriant; 
Kochia sedifolia, Atriplex vesicarium, and others occur some- 
| what closer together. The annual flora resembles that’ of the 
plains themselves, but certain perennial plants appear in this 
zone, e.g., Solanum. esuriale, S. coactiliferum, Sida corru- 
gata. On disturbed ground near the village Calandrinia 
polyandra and Euphorbia Drummondii are found in some 
quantity. 


. 


548 


Sandhills —The vegetation of the sandhills to the east. 
of Ooldea is a marked contrast to that of the Nullarbor Plain. 
In place of the dwarf grey or white Chenopodiaceous bushes 
of the latter, the sandhills bear a relatively luxuriant covering 
of trees and shrubs. These differ both in their much larger 
size; and also very much in leaf form. While the plants on — 
the plain have more or less spreading, though small, leaves — 
which are succulent and covered with hairs, almost all the 
plants on the sandhills have smooth leaves which are hard im 
texture and placed with their edges to the light, being either — 
pendant or vertical. The leaves are either quite glabrous and — 
polished as in Hucalyptus, spp., and Myoporum, or grey in 
colour, due either to wax or to a covering of very small hairs” 
that do not spread from the surface, as in several species” 
of Acacia. Even those plants such as Casuarina, Bossiaea, — 
and others which are almost or quite leafless have their assim-— 
ilatory branches erect or pendant, not spreading. This leaf | 
character applies both to the larger bushes and trees,’ and 
also to the smaller undershrubs which here bear, for. the ‘most 
part, small hard leaves placed more or less vertically. ‘This 
difference in leaf type makes the vegetation on the two parts” 
very distinct, even when seen from long distances (pls. xxxill. 
RKIVe ae, se eunelixoey ge?! Tri both situations it may be. : 
remarked that all the plants are evergreen; not a single 
deciduous plant was found. 


For purposes of description the vegetation can be divided 
into three portions: (1) the sand ridges, (2) the hollows 
between, and (3) the basin known as Ooldea Soak. Even with 
this division the vegetation presents at first glance a rather 
bewildering lack of “uniformity ; many plants are apparently 
localized in their distribution, and’ situations externally very ~ 
similar often bear different plant populations. This variability | 
can to some extent be explained by a recognition of the fact | 
that the sand is not uniformly stabilized. The plants in | 
different places vary in their efficiency as sand retainers. It | 
may also be correlated with’ the frequent limitation of areas — 
‘drenched by rainstorms. These ‘‘patchy’’ rainfalls may, make — 
certain places good seedbeds, while the surrounding aréas are 
too dry for a high percentage germination or even any at all | 
that season. The result is that Grhine some sandhills are ablaze | 
with the flowers of an abundant annual flora, others a few 
miles away are without any appreciable annual growth at all. 
It is well known to pastoralists that the season at which'a | 
soaking rain. falls profoundly affects the type of annual flora | 
that results. It seems to us. quite legitimate to assume that 
the germination of other plants is affected also, and hence that 
different phases of an open flora may be shown under similar 


ee 


£ 
=e 
we 


549 


edaphic conditions within a short distance of each other, 
because the climatic factor of rainfall has varied as a result 
of some fortuitous circumstances, e.g., a local thunderstorm. 
A further factor affecting this variability is interference by 
man, which has been not inconsiderable in some places. This 
interference is partly caused by the aborigines, who cut down 
and uproot trees and shrubs around their camps in an aston- 
ishingly reckless manner, and partly to the demand for wood 
during the construction of the railway line. 

The sand ridges have a rather varied flora, among the 
most prominent and generally distributed plants on the ridges 
are Acacia linophylla, known as the ‘‘sandhill mulga’”’ ; A. lagu- 
lata, Dodonaea attenuata, and, rather less generally distri- 
buted, Leptospermum laevigatum, var. minus. These, with 
locally some quantity of Hakea leucoptera, Grevillea steno- 


botrya, Grevillea guncifolia, and Bossiaea Walkert, mark the 


earlier stages in the stabilization of the sand. Frequently 
communities of these plants occupy the crest of the ridge while 
the sides of it have others which represent the result of a 
more stable condition. The most prominent. of these are 
mallee forms of Hucalyptus. The most common are /. oleosa, 
EL. leptophylla, and FE. sp. affin. oleosa, while EF. transconti- 
nentalis is rather more local. Other plants here are Acacia 
Randelliana, A. Oswaldii, A. aneura, EHremophila alterm- 
fola, and Cassia eremophila.. In some parts Callitris verru- 
cosa, Grevillea netatophylla, and Hakea multilineata occur in 
this situation. These last three species were all seen at 
Immarna, 20 miles east of Ooldea, but not in the immediate 
vicinity. 

On some of the sand ridges near the margin of the plain 
Casuarina lepidophloia occurred both on the sides and even 
extending on to the crests of the ridge, but this was not 
general. When the sandhills had become more stabilized the 
mallee and its associated plants extended over almost the 
whole, occupying both sides and crests. Indeed, the notable 
Hucalyptus pyriformis seemed only on crests of ridges. 
Whether the covering consists of Acacias and Leptospermum 
or mallees the canopy is not continuous, considerable spaces 
being left between most of the plants. A marked feature, 
and cne most obvious with the plants on the crests, is the 
presence of large quantities of dead branches and wood. 

Beneath the trees and bushes a moderate amount of 
undergrowth occurred in some places, though in parts the 


_ sand was practically bare. Of perennials, on the crests there 


occurred Pimelea microcephala, Rhagodia Preissii, while on 
the slopes the ‘‘porcupine grass’’ Triodia irritans was locally 
very abundant, forming what appeared at a distance to be a 
continuous cover; associated with it was Bassia echinopsila. 


550 


Locally, -annuals were abundant on these sand ridges; 
these were largely composites. The annual flora is strikingly 
different from that occurring on the plain. On the crests of 
the ridges, and apparently confined to such situations, were 
Calandrima disperma, Stenopetalum lineare, Myriocephalus 
thisocephalus, Waitza acuminata, and Podotheca angustifolia. 

Others are less restricted in their habitat, as Triglo- 
chin centrocarpa, Trichinvum alopecuroideum, Calandrinia 
volubilis, Stenopetalum sphaerocarpum, Crassula verticillaris, 
Erodium cygnorum (very local), Zygophyllum sp., Poranthera 
microphylla, Didiscus cyanopetalus, Minuria leptophylla 
(perennial), Brachycome ciliaris, Angianthus tomentosus, 
Helichrysum ambiguum (perennial), H. Lawrencella, H. 
floribundum, H. hyalospermum, H. strictum, H. moschatum, 
H. rosewm, Senecio Gregoru, S. brachyglossus. 

The geophyte Thysanotus exiliflorus also occurs) here. These 
herbaceous plants for the most part grow on the sand in the 
spaces between the bushes and not immediately under them. 
Below most of the bushes or trees there was an area covered 
by dead leaves, fruit, branches, etc. This appeared to pre- 
vent the development of an annual flora. 


Hollows between Sandhills —While in a general way a 
distinct flora for sandhill hollows can be recognized, this flora 
varies greatly in accordance with the amount of sand that is 
present in the hollow. Im nearly all cases the soil is much 
firmer in the hollows than on the ridges and the vegetation 
less dense and more easily penetrated, largely owing to the 
presence of tallish trees in addition to the bushes. The 
generally most abundant plants are Myoporum platycarpum, 
which is a tree 20 to 30 ft. in height, with Heterodendron 
oleifolium, a bush or small tree. These form the general 
character plants, but grow in association with many others. 
Of trees Pittosporum phillyraeowdes is most abundant, and 
of bushes Acacia Randelliana, A. aneura, A. Oswaldu, A. 
collettoides, Eremophila glabra, EF. Latrobei, LE. altermfoha, 
Cassia Sturt, C. eremophila, Fusanus acuminatus, F. persi- 
carius, Dodonaea microzyga. When more sand is present 
Casuarina lemdophloia becomes abundant, growing into large 
trees, which form an open forest, which will be referred to 
later. As in all the communities around Ooldea the plants 
are rarely in close contact with one another, but stand at 
intervals. 

The undergrowth in the hollows is variable both m 
amount and in composition. In parte it is quite absent, but 
for the most part some undershrubs or herbs are present. Of 
the former QOlearra Mueller and Westringia rigida are the 


551 


most abundant, together with Vetraria Schoeberi and Zygo- 
phyllum fruticulosum, which are more local. 

In some parts, and more especially towards the margin of 
the Nullarbor Plain, where very little sand is present in the 
hollows, Atriplex vesicarvum and other species may occur in 
some quantity ; also Cratystylis conocephala is abundant. This 
last is a plant which at first sight bears a most striking 
resemblance to Kochia sedifolia, a resemblance that is empha- 
sized when it is growing with Atriplex spp. (cf. Tate, loc. 
cit.). Other perennials in the undergrowth are Khagodia 
spinescens, var. deltophylla, and Scaevola synescens. 

The annual herbaceous flora is much poorer than that of 
the ridges, and but few species appear limited to this habitat, 
é.g., Calotis hispidula, Brachycome pachyptera, and 
Trichinium imecanum. Some species, however, are more 
abundant here, as Helipterum strictum, Tetragoma. expansa, 
and Pimelea swmplex; Stipa, sp., apparently dead, was also 
locally abundant, and Danthonia penicillata occurred occasion- 
ally. When the limestone soil came to the surface in a hollow’ 
the annuals of the plain were present. 


“Oak’’ Forest.—Forests of Casuarina leydophloia occur 
to the south of Ooldea. In this vicinity the forests have been 
much reduced in quantity owing to the utilization of the 
timber for condensers and other activities associated with the 
construction of the railway. At present untouched forest is 
not met till about seven miles are traversed (pl. xxxiv., fig. 2). 


The forest occupies rather flat hollows between sand 
ridges. The soil, however, is sandy and rather loose all 


through, even in the centre of the flats. The Casuarinas 


extend on to the sides of the surrounding ridges but not on to 
the crests of them, which are covered by Acacia linophylla, 
A. aneuvra, and A. Oswaldiu, i.e., with a typical sandhill 
crest community. | 

The forest is a very open one, and the trees are often of 
considerable size. Measurements of some of them showed a 
diameter of 26 in. at 1 ft. from the ground, and a height 
of approximately 50 ft. was estimated. When mature the 
trees have spreading branches, though in the young condi- 
tion their habit is somewhat strict and pyramidal. * Numerous 
young trees were coming up in the forest. The ‘‘oaks’’ far 
overtopped any other plants, the other trees present being 
much smaller; these are Myoporum platycarpum and mallees 
(EL. oleosa, FE. sp. affin. oleosa, E. leptophylla). The 
latter occur on the slopes and ridges, and may reach a height 
as great as Myoporum, i.e., 20 ft. to 30 ft.: Below, and 
especially between the trees which nowhere form a continuous 


552 


canopy, is a considerable assemblage of shrubs or bushes. Of 
the most prominent are /wsanus acuminatus, I. persicarius, 
Dodonaea attenuata, D. microzyga, Hetereodendrow olei- 
folium, Acacia Randelliana, A. colletioides, A. tetragono- 
phylla, A. Oswaldu, A. linophylla (locally), Cassia Sturti, 
C. eremophila, Eremophila Latrobe, E. alternifolia. As 
undershrubs there occurred Olearia Mueller, Westrimgia 
rigida, and Atriplex vesicarvwm, all, however, rather local. 
Annuals and herbaceous plants were not at all prominent, a 
few individuals only being noticed of Zygophyllum ovatum, 
Calandrinia volubtlhs, and Helipterum floribundum. 

While the forest of Casuarina lemdophloia is rather 
limited in its distribution around Ooldea, further to the east, 
as can be seen from the railway, it commonly occupies the 
sandy hollows, while mallee occurs on the ridges. 

The part where the Casuarina trees have been cut down 
shows some signs of regeneration; young trees are springing 
up in places in some quantity. The other plants of the forest 
- have been left by the timber-getters, and in the cut areas form 
an open scrub with many more undershrubs than occur in 
the forest itself. Atriplex vesicarium is abundant in parts, 
and in others Olearra Muellert and Cratystylis conocephala 
form a distinct layer of undershrubs. Stipa setacea is also 
abundant locally. Annuals, too, are more frequent here, 
though not very prominent. Besides those of the forest there 
were Helipterum strictum, Helichrysum Lawrencella, Angi- 
anthus tomentosus, Salsola kali, and Lepidium phlebopetalum. 


Ooldea Soak.—It will be recalled that the Séak is a basin 
in the sandhills, itself filled with sand that is thrown into 
small ridges. This sand is white in colour, not red, and is 
rather looser in texture than that in other parts. On the 
sandhills composed of this white sand in the part just 
around the Soak certain differences occur in the plant popula-— 
tion; Acacia linophylla is rather less abundant and Lepto- 
spermum laevigatum more so. Also a certain number of plants 
grow on this white sand which were not noticed elsewhere. 
Among these may be mentioned Grevillea stenobotrya, Hakea 
leucoptera,-Gyrostemon ramulosus, and Hucalyptus pyriformis. 

The basin of the Soak itself has been the seat of some 
interference. owing to man’s activities. For centuries this 
has been a camping ground for aborigines, and in more recent 
times the white man has utilized it. As a result, a consider- 
able part of the basin is bare sand, which is liable to drift, 
and which has only the tops of the ridges occupied by plants. ~ 
The most abundant are Leptospermwm laevigatum, var. 
minus, and Melaleuca parviflora. Other plants are not 


553 


prominent; indeed, one or both of these two are often the only 
plants present. The other species present are Dodonaca 
attenuata, Acacia ligulata, Cassia eremophila, with locally 
Hibbertia crispula and Adriana tomentosa, the lastnamed in 
hollows. On the bare sand there were occasional plants of 
Salsola kali, Myriocephalus Stuartu, and a few tufts of Stzpa, 
sp. These scattered tufts appeared dead and were much eaten 
down by rabbits. 

In the Soak the water-table is about 5 ft. to 15 ft. below 
the surface, but yet this seems to have very little effect on the 
plants. Adriana tomentosa, and possibly Melaleuca, are the 
only species that seem dependent on the presence of ground 
water. Adriana, which is known as the ‘‘water bush,’’ has 
a leaf very different from any other plant found in the district. 
It is relatively thin and spreading horizontally, neither at all 
succulent nor hard and coriaceous. The leaves and shoots of 
this plant have a much smaller water content than most of the 
other plants found. 

Leptospermum, which is such a feature of the sandhills 
in the basin, also occurs on many of the sandhill ridges in 
positions quite remote from supplies of bottom water, and it 
seems better regarded as a plant characteristic of loose sand. 
than of moisture. 


Salt Lake—Separated from the Soak by a high ridge of 
sand with the typical vegetation of Acaczas, etc., is another 
basin occupied by a small salt lake (pl. xxxvi., fig. 1). At 
the time of our visit this lake was dry, and the loose level soil 
of the bed was largely composed of soluble crystals, amongst 
which gypsum crystals were numerous. 

The main bed of this lake was bare of plants, but round 
the margins occurred a halophytic vegetation. That nearest 
to the bed was an open community of Arthrocnemum, spp. 
(A. halocnemoides, A. halocnemoides, var. pergranulatum, 
and Arthrocnemum, sp.). At a slightly higher level occurred 
Frankema fruticulosa, often almost pure or mixed with a few 
plants of Atriplex paludesum, Bassia diacantha, Kochia brevi- 
folia, Mesembryanthemum aequilaterale, Nitraria Schoeberi, 
while still higher on the sand just below the typical bushes of 
the surrounding sandhill flora occurred Salsola kali, Atriplex 
vesicuria, Stipa, sp. (dead tufts), Calandrinia volubilis, and 
Didiscus cyawopetalus. Beyond this zone one passed into the 
typical sandhill flora (pl. xxxvi., fig. 2).. 


RELATIONSHIPS OF VEGETATION. 
The salt lake just described was interesting, especi- 
ally as demonstrating the total change of flora that 


q 


occurs in those parts where the soluble salts in the soil 
reach a high concentration. None of the species of the 
‘‘saltbush’’ or “‘bluebush’’ vegetation of the plains. which 
have so often been termed halophytes by other workers, were 
present around the lake, with the exception of A triplex vesi- 
carium in small quantities near the extreme margin. The | 
Atriplex of the salt lake was Atriplex paludosum, which also 
occurs in coastal salt swamps; it 1s a true halophyte. The fact 
that halophytic vegetation 1s composed of members of the 
Chenopodiaceae, and often of species of the same genera that 
occur on the Nullarbor Plain, cannot be taken as proving that 
the character plants of the latter habitat are halophytes, as 
was assumed by Cannon. This distinction between the halo- — 
phytes of a salt lake and the ‘‘saltbush’’ or ‘‘bluebush’’ com- 
munities has previously been noted by Tate (/oc. cit.). We 
regard it as an important one. 

In any climate where the evaporation rate is in excess of 
the precipitation the soil will tend to accumulate considerable 
quantities of soluble salts. But in no part of the Nullarbor 
Plain was there any sign of surface accumulation of crystals. 

The communities of Kochia and Atriplex that occupy the 
- surface of the Nullarbor Plain with their rich crop of annuals 
ought to be regarded as representing a semi-desert flora rather 
than a halophytic one. The bluebush, and to a less extent 
the saltbush, seem to represent the succulent flora for this — 
region. Comparisons would be better made between them 
and the constituents of the succulent communities of South 
Africa or America than with halophytic plants. 

On the Nullarbor Plain the vegetation of the dongas, with 
the scattered small trees and bushes, represents a further 
advance that takes place in that area with increasing amounts 
of soil, and especially of moisture. The communities of the 
dongas were surprisingly like those developed on the sandhill 
hollows. They appear to exist under very similar conditions, 
though the vegetation is much more sparse and less advanced 
owing to the more severe environmental conditions. Con- — 
sidered in relation to the plain formation as a whole these com- 
munities of plants in the dongas appear to represent what | 
Clements (5) would term a post-climax: that is to say, while 

| 
| 
| 


054 


4 


tps 


the general climatic and edaphic conditions cause a stoppage __ 
of development on the plain at the stage of an open com- 
munity of bluebush, in the slightly more favourable conditions ~ 
in the dongas the process is carried to a further stage. This 
further development can be traced to some extent; in the — 
smallest depressions only very xerophyllous species occur, €.g., 
Acacia tetragonophylla; larger depressions with more soil have 


(15) Plant Succession, 1916, p. 109. 


5DD 


Acacia aneura, and sometimes Pittosporum phillyracoides; 
while in the largest, which have most soil, the furthest devel- 
opment occurs, and Casuarina lepidophloia forms sniall scraps 
ef woodland. 

The vegetation of the sandhills stands in marked contrast 
to that of the Nullarbor Plain. Under the same climatic con- 
ditions on the sand, even in the most exposed positions, the 
plant communities are of large woody plants, and the general 
effect is of some luxuriance. This effect is perhaps more 
apparent than real, but is certainly marked as compared with 
the plain. The difference must be attributed to the difference 
in soil. The loose sand has practically no run-off—all the rain 
falling percolates into the soil at once. Further, the sand 
readily forms a quite dry dust mulch on the surface which 
prevents loss by evaporation, whilst the relatively coarse soil 


_ particles carry on a certain amount of condensation. The 


much finer-grained soil on the plain, on the other hand, will 
not condense, and, owing to its much greater water-raising 
power, will lose water by evaporation instead of forming a 
mulch. As was pointed out above, on the sand we have two 
distinct sets of communities—those in the hollows and those 
on the ridges. The former bear a close relation to the com- 
munities existing in the dongas, but development proceeds 


-further owing to the better soil and-the shelter afforded by 
_ the ridges. 


Two moderately distinct communities can be recognized, 
namely, that of Myoporum platycarpum and Heterodendron 


_ olerfolium and the open forest of Casuarina legdophloia. The 


former occurs where less sand is present. In both cases some 
developmental stages can be recognized, especially when one 
compares some of the hollows near the margin of the plain 
with those further east. The early stages are represented by 
Atriples, vesicartwm generally with bushes, especially of 
Acacia anuera, A. Randelliana, and Hremophila Latrobe:. In 
the Myoporum-Heterodendron phase, which appears as a 
climax, the Atriplex disappears, but the other bushes are still 
present. When more sand is present the climax appears to 
be the Casuarina lepidophloia forest. It is noticeable that 
this, rather than the Myoporum-Heterodendron community, 
occupies most of the hollows further east, where the sand has 
become more distributed. The portions of this forest which 
have been cut down show a return to the earlier phase with 
considerable quantities of Atriplex vesicarium as under- 
growth. 

The sandhill ridges, as described earlier, exhibit a series 


_ of developmental phases of which mallee (/ucalyptus) 


appears to be the climax. Here again the succession generally 


556 


reaches a more advanced phase as one passes east for 15 to 
20 miles, where mallee is found to cover almost all the sand 
ridges. On the other hand, nearer the plains, the presumably 
younger sandhills are almost or quite without mallee and are 
occupied by Acacias. 

To summarize, excluding the salt lake with its highly — 
saline special conditions, one can recognize probably four types 
of vegetation :—/a@) The Nuilarbor Plain, with its open .com- 
munities of Aochia sedifolia or Atriplex spp.; (6) the sand- 
hill ridges commencing with Leptospermum laevigatum and 
Acacia linophylla, etc., and culminating in an open mallee 
commuuity; (¢) the sandhill hollows with sandy soil culmin-— 
ating in open forest of Casuarinu lepidophloia; and lastly, 
(d) the hollows with firmer soil which appear to reach a climax > 
in the community of IJyoporum platycarpum and Hetero- 
dendron oleifolium. These last two are very closely allied and 
all sorts of transitions with intermediate conditions can be 
noted. 

The donga communities represent attempts stopped by 
conditions to develop upon the open plain the vegetation char- 
acteristic of one or other of these last two types. 


FLORA. 


A list of the flora so far as we have collected it is given 
below (Appendix). This has been enlarged by the inclusion” 
of thirty species recorded or collected ‘by Black from the 
Ooldea district, but not seen by us. The total number of 
species amounts to 188. 

In this list we have given the habitats of the, plants 
dividing the district into six main divisions, viz., the Nullarbor 
Plain, the dongas on the plain, the sand ridges, the sand flats 
between the ridges, the Soak, and the salt lake. From such 
a list the several plant communities recognized by us can easily 
be seen to have their characteristic floras. Necessarily the 
classification is somewhat arbitrary, special difficulty being 
found in deciding whether a plant growing at the edge of the 
Nullarbor Plain properly belongs to the plain flora or to that 
of the sandhill areas. Nevertheless, it becomes clear that, as 
we have stated above, the donga flora has more resemblance to 
that of the sandhill country than to that of the plain. 

Secondly, it will be seen that in the sandhill area the floras 
of ridges and of flats are strikingly distinct, sufficiently so 
recognize them as different associations. Again, however, it has 
not been possible to distinguish in tabular form between those 
flats with a deep sandy or loamy soil, and those in which the 
underlying limestone comes near to the surface. These last 
are recognized as being inliers of the plain association that 


557 


have become included in the sandhill area, though in the list 
they are included as sandhill flats. 


Finally, in the list, the plants are classified according to 
their ‘‘life-form,’’ using Raunkiaer’s system.4) As no such 
examination of an Australian flora has been made before, the 
ten life-forms recognized by Raunkiaer are briefly defined 
below. 

-PHANAEROPHYTES are plants whose dormant buds project 
freely into the air, ¢.e., trees and shrubs. They are com- 
monly subdivided according to their height into four 
groups :— 

- Megaphanaerophytes, tall trees, over 30 m. 
Mesophanaerophytes, medium-sized trees, 8-30 m. MM. 
_ Microphanaerophytes, small trees and shrubs, 2-8 m. 
Symbol M,. 
Nanophanaerophytes, shrubs, 2 m. and less. Symbol N. 


‘e- CHAMAEPHYTES (@hh.) are plants with buds or shoot 
apices perennating on the surface of the ground or just above 
| it (under 25 cm.). These buds gain some protection either 
| by snow, or, in dry countries, by dead plant remains. 
| .  HemicryptopHytes (fl.) have their dormant buds in 
: the upper soil crust, just below the surface, thereby gaining 
additional protection. The aerial parts are herbaceous and 
die away at the onset of the critical period. 


_ CrypropHytTes are plants with their dormant parts well 
buried in the case of geophytes (G.), the only subdivision of 
the class present. in the Ooldea flora. Marsh plants (helo- 
phytes) such as Typha and Phragmites, and some aquatic 
plants (hydrophytes) as. Nymphaea and Potamogeton form 
the other subdivision (HH. “) which is not represented at 
Ooldea. 


THEROPHYTES (Tin. ) are plants the seeds of which germ- 
inate rapidly at the favourable season, soon pass into flower and 
fruit, and then die away. These, therefore, pass the unfavour- 
able season as seeds. They are ‘all annuals, and many in the 
floras of arid regions are ephemeral. Two other classes are 
Stem SuccuLEents, notably scarce in the flora of Australia as 
a whole, and EPIPHYTES (E.). Strictly speaking, these 
perched plants should gain nothing but an elevated position 
from the phanaerophytes on which they grow, hence they are 
best developed-in wet regions. No true epiphytes occur at 


. @4)-Smith, W:* G., Raunkiaer’s Life-forms and Statistical 
Methods, Journ. Ecol., i:, pp. 16-26, 1913; in this paper the 
literature is summarized to date. Taylor, W., Growth- forms of 
the Flora of New. York ae Vicinity. ; Am. Jo ourn, , Bot., 1. . pp. 
23-31, 1915. , 


558 


Ooldea, or indeed in South Australia, but we have placed 
in this class the many parasites, chiefly Loranthus, spp., which 
are found in the Ooldea florula. 

The classification of a flora into these life-forms may be 
utilized in the following way, as illustrated by the table below 
(Table III.). For any locality the total number of species 
analysed is given, followed by the results given as percent- 
ages according to the grouping above. ‘‘Such an analysis for 
any region is termed the biological or phyto-climatic spectrum. 
The normal spectrum is the base line, and the outstanding 
features of the other spectra are deduced by comparison, not 
by the highest: percentage in their own curve, but by the 
amount of variation from the normal spectrum. The latter is 
ideally the phyto-climatic spectrum of the whole earth; 
actually it is obtained by computation, and at present is given — 
only as approximate. It was arrived at by first selecting 
1,000 representative species and then taking 400 of these, 
which were carefully analysed. This number, 400, has been 
carefully controlled in various ways,’’ 5) which, however, need 
not be considered here. 

: TasLeE III. 

Biological spectrum of Ooldea district compared with that of 
other arid regions :— 

Percentage of Species belonging to each Life-form 


Total ha . 


speciesMM M N Ch H G HH TH E S 
Normal Spectrum 400 6 17 20 9 27 3 L 5 Wiis 
Ooldea ... . 388 5 19 283 14 4 5 — 85 4° — 
Libyan Desert ... 194 — 3 9 24 20 4 1 42 — — 
Aden ... . 176 — 126 27 19 3 =| a= a8 
Madeira Lowlands 213 — 114 7 24 — 3 SI — — 

ee ee 

Transcaspian ... 768 10 7279 5 4 — — 
Death Valley ... 294 — 221 7 18 2 >» 4 — $ 


* These epiphytes are all parasites, not true epiphytes, see text. 


Considering the spectrum of the Ooldea region in relation 
to the normal, the absence of tall trees,“ (MIM.) is as 
marked a feature as is the smaller number of perennial 
herbaceous plants, 2.e., hemicryptophytes and geophytes. On 
the other hand, the micro and nanophanerophytes, and also 
the chamaephytes, are all in excess of the normal percentage. 
The departure from the normal is most marked in the chamae- 
phytes (5 per cent. increase), but the two Phanaerophyte 


(15) Smith, W. G., loc. cit., p. 18. 
(16) Casuarina lepidophloia, as found growing at the ‘Oak 
forest,’ was the only tree over 8 metres. 


559 

groups M. and N. each show 3 per cent. above the normal. 
This indicates that the environmental factor favours low trees 
or shrubs. The most marked departure from the normal is 
in the therophytes. Annuals are a very prominent feature 
in the Ooldea florula as far as it is known at present, while 
it is probable that collections made in a “‘good season’’ would 
still further enlarge the class. 

These deviations from the normal become the more inter- 
esting when considered in relation to the biologic spectra of 
other arid regions. Closest correspondence is seen between 
that of Ooldea and the Death Valley, California. The thero- 
phyte percentage of 34 is sufficient to mark the Ooldea region 
as belonging to the desert series, but the number of low 
woody plants is exceptional. The micro- and nanophanaero- 
phytes together (Ml. and N.) amount to 43 per cent., a 
number in excess of both the normal spectrum or that of any 
other arid region available for comparison. Further work on 
the Australian flora is needed before the significance of this 
can be fully appreciated. While the Ooldea region may be 
said to show a therophyte flora, the tendency of its peren- 
nials to be woody plants and to have their resting buds above 
the surface of the ground and not below it (Class Ad. is re- 
markably subnormal) is certainly a point that calls for further 

, investigation. 


Department of Botany, University of Adelaide. 


APPENDIX. 


The following list contains the complete flora of Ooldea so 
far as recorded at present. We have distinguished the species 
recorded by Black and not seen by us by prefixing the letter B., 
and two species of Acacia are given on Cannon’s authority. The 
six divisions showing the habitat are shown by the following con- 
tractions:—Nullarbor Plain, N.P.; Dongas, D.; Sand ridges, 
S.R.; Sand flats, S.F.; Soak, S.; and Salt Lake, S.L. 


a. 
aa (M) lela) fa 
) Bo [2 [Ala loi jo | a 
A ee Coe eee | 
| Callitris verrucosa, R. Br... MM. flop eet 
Triglochin centrocarpa, Hook. Th. | cae came 
B.| Panicum gracile, R. Br ... BL. Ll eae i | 
B.| Amphipogon strictus, R. Br., v. - gracilis | 15 | i+i+| | 
| Stipa setacea, R. Br. ben ite 4 H. |+/+| | [+/+ 
S. seabra, Lindl. Bore || 
B.|S. sclerata, Behr. H. | iH | 
Danthonia ’penicillata, F. v. M. H. |+/°; |+ | 
imiodiasyeritans, R.Br. os. 2X: “us Ch. \-+| | 


560 


| ea } 
ao lai 
=O e 
tie 
| | | ae 
Lomandra leucocephala, (R. Br.) Ewart aH Ch.) i+ | | | 
Thysanotus exiliflorus, F. v. M. ... G. | +| | 
| Casuarina lepidophloia, F: vy. M. ... MM.| |+/+/+! | 
| Grevillea juncifolia, Hook. M. | +| | | 
G. stenobotrya, F. v. M. ... sil M. | | +) \* 
G. nematophylla, F. v. M. al nes +] 
| Hakea leucoptera, R. Br. 2 | a +} k 
H. multilineata, Meiss. ... : M. |}. + | 
Fusanus acuminatus, R. Br. M. +| & 
F. persicarius, F. v. M N: + + 
B.| F. spicatus, He Biro M. Ls 
Viscum articulatum, Burm. EK. +| | 
Loranthus exocarpi, Behr. E. i+ 
L. Preissii, Mig. K. | ia | 
L. quandang, Lindl. E. ye +} | | 
B.| L. miraculosus, Mig. E. [s  aeicet 
Be a pendulus, Sieber . EB. | \5ea ee 
. Miquelii, Lehm. ... Bi] eee 
Rhasodia spinescens, R. Br., var. delto- | Gy 
| phylla Paes _ N. |+] | Zh 
R. Gaudichaudiana, Moo. No is 
B.| R. Billardierii, Moa. se Ne ent A ie Fate: 
R. Preissii, Mog. a ess ei | I] | 
Chenopodium micr ophyllum, F. v. M. | Ka | 
var. desertorum, J. M. Bo Phd laa bs | | |4- 
C. nitraraceum, F. v. M. hora |+- 
Atriplex vesiearium, Hew. | ON. [sey Se 
| A. holocarpum, F. | Th, |e) ee 
| A. paludosum, R. Be en OR | | 
| A. stipitatum, Benth. .| Ch. |. eae 
ANOS Sy Peter aa Ch. |+| | | | 
Kochia appressa, ‘Benth. ey frre Re ie ee) 
| K. brevifolia, R. Br. jee) | ieee 
FG pyramidata, Benth. : N. |+/ | J4I 
| K. sedifolia, F. v. M. N. |+{ | [+] 
| K. triptera, var. erioclada, Benth. , RS i+ 
K. aphylla, (Ree | ON. [+ | 
Bassia scleroloenioides, ‘F. v. M. | Ch. |+i/4+] i+ 
|B. diacantha, F. v. ..| Ch. [+/+] J+] 
| B. echinopsila, F. v. M. ... Sahonehie. 4 |+| 
| Enchylaena tomentosa, R. Bri wuic, i aN | | 
| Arthrocnemum halocnemoides, Nees. [ae ae | | 
te halocnemoides, var. cena _ | | ) 
| A., fi ‘ib aih| | ae 
lSateots, kali, Le Ss _ strobilifera, “Benth. a! ide i + it 
Trichinium incanum, R. Br. Pee a ye | | |+| 
T. incanum, vy. erandiflorum, ‘Benth. ses | 
Lek; alopecuroideum, Tyas. PeThes | 1 sel 
| T. alopecuroideum, v. rubriflorum, J.M.B.| Th. ey |+ 
| Gyrostemum ramulosus, Desf. xis aM, | | {+] \* 
| Tetragonia expansa, Murray ... ... ...| Th. |+| |+ 


561 


C | 


Bs. 


B. 


B.| Beveria opaca, F. v. M. 


.|C. pusilla, Lindl. 


| Acacia colletioides, A. 


Me i Reade a aequilaterale, Haw. 
era. volubilis, ‘Benth. 

C. disperma, J. 

C. polyandra, (Haoks\ Benth. 


ke 


Cassytha melantha, R. Br. 

Alyssum minimum, Pallas 

| Stenopetalum lineare, at Br. 

S. sphaerocarpum, F. MES. 2: 

| Lepidium phils Miopakakcas, F. v. M. 
| L. Draba, L. é 
Crassula verticiliaris, DC... 
| Pittosporum phillyraeoides, DC. 
Cunn. 

M. 


{ 
| 
| 
| 
E 
| 
| 
| 
| 
/ 


| A A. tetragonophylla, F. v. 
| A. ligulata, A. Cunn. 
| A. Oswaldii, F. v. M. 
A. Randelliana, Ww. Y. Pitz. 
| A. aneura, F. v. M. 
on aneura, v. latifolia, J, EB. 
| A. linophylla, W. V. Fitzg. 
| A. brachystachya, Benth. 
| A. Kemplana, aie. Mic 
i Gassya Sturtii, i. Br. 
-C. eremophila, A. Cunn. - 
/C. eremophila, v. zygophylla 
/C. eremophila, v. piaspods ae: 
| C. phyllodinia, Re Br: as 
Swainsonia colutoides, F. v. M. 
Bossiaea Walkeri, F. v. ee 
| Lotus australis, Andrews, 
| EKrodium cygnorum, Nees : 
| Zs gophyllum fruticulosum, DC. 
|Z. ovatum . 
| Z. ammophilum, F. 
Pee 43 
| Nitraria Schoeberi, Linn. 2 
Boronia coerulescens, Pty MM. 
| Euphorbia Drummondii, Boiss. 
| E. eremophila, A. Cunn. 
| Adriana tomentosa, Gaudich. 


_M. 


| Poranthera microphylla, Broug. 
| Stackhousia viminea, Smith. 

| Heterodendron oleifolium, Desf. 
| Dedonaea attenuata, A. Cunn. 
| D. microzyga, F. v. M. 
'Lavatera plebeja, Sims, 
| Hook. 

| Sida corrugata, 
|S. petrophila, F. v. 


var. 


Lindl. 
\.': Bees 


_ pubescens 


tomentosa, 


Raunkiaer 
Class 


562 


ie. | ee 
| 3 lol tele fe 
| 3° |e ed v5 
ones) Les lcuoetiae 
Hibbertia‘crispula, J. iM Gee) ee a | |+ 
Frankenia fruticulosa, DC. ....... ...| Ch. 4 
B.| Lythrum hyssopifolia, L. ah A er ne wee i | ae 
Pimelia microcephala, R. Br. IN: +/ | a 
P. simplex, F. v. M. ; Th. + |] 4 
| Eucalyptus pyriformis, Turez. M. Hy i 4 
E. leptophylla, Mig. . M. +] | 
E. oleosa, F. Cece Ei tee M. +! |. | 
B.,.sp.. indet. affin. oleosa ... .. | M. a | 
E. transcontinentalis, J. H. M. .. M. +} | | = F. 
EK. Pimpiniana, J. H. M. PAD ioe ty ( ro a en +} | | 3 
maybe ethic eee v. minus,} M. +| [+ | . 
Melaleuca parviflora, “Lindl. N. | , i 
Didiscus cyanopetalus, Psy Me Th. + | | - | 
B.| Alyxia buxifolia, R. Br. ... | NE +} . 
Convolulus erubescens, Sims. Th. + ce 7 
B.| Heliotropum europeum, L. | Cher! Leake oie 
B.| Halgania cyanea, Lindl. He Ch. | ee | | 
Echinospermum concavum, F. v. M. Th. [+/+] |+| | | 
B.| Dicrastylis beveridgei, F. v. M. N. | ae | 
Westringia rigidia., R. Br. N. ; |+l | 
B.| W. Dampieri, R. Br. | cae + | 7 
Solanum esuriale, Lindl. | Ch. |+ | | a 
S. coactiliferum, FMB: | Ch. |+ i+{ | |, 
B.| S. hystrix, R. Br. Ch. | . 
Nicotiana suavoleus, Lehm. 1 2h. +1 fj | 7 
Myoporum platycarpum, R. Br. ... whe Ios SO eA . a 
Eremophila oppositifolia, BBE 6 he he Ft 
E. Latrobii, F. v. M. E re ee he ~ F 
b.| E. Latrobia, v. Tietkensii, ‘J. M. B. Reel! EU | | | 5 
E. maculata, bv, Mie aes | M. | | . 
B.| E. Paisleyi, BF. v. M. | M. hoot i 
B.| E. Goodwini, F. v. M. ... [ a EA eae be | | 
BE. glabra, (R. Br.) Ostenf. | M. | aes 
E. alternifolia, A cus Tek. at M. | +)/+i+ | | 
| E. longifolia, F. v. M. | M.). Tee | a i 
B.| Pholidia scoparia, F. v. M. | M. | lis eds | 
| Plantago varia, R. Br. | Th. | +/+] ‘ 
| Pomax umbellata, OME cc) Pane: | Th. + | | § 
B.| Goodenia strophiolata, yo: | Ch. +/+ 3; 
|G. ‘pusillifiora; a ML, 63.) | Th. |+ | ‘ 
|G. pinnatifida, Schl. vee | Th. |+/+{) (+ . 
B.| Dampiera lanceolata, A. Cunn. Pe. ak | . 
| Scaevola spinescens, R. Br. EP | i+} | , 
‘| Olearia Muelleri, Benth. N. | | | ae 2 4 
| O.s subspicata, Benth. N. | | be | 
| Minuria leptophylla, DC. CE i 5 
| Calotis hispidula, F. v. M. ee + e 
B.| C. erinacea, Steetz ... ws | Th it te | 
| Brachycome pachy aio F. v. M. The 4) 3) 
|B. ciliaris, Lees. hy | Th.) dds 


Lc 


563 
: 2a 
| EE A} lei) le 
| S ZlIA|Ni vin \|n 
jet 
| 
| Cratystylis conocephala, S. Moore. ...|_N. | | | he : 
| Iscetopsis graminifolia, Turcz.  . “ Th. Pea i+} | 
| Myriocephalus rhizocephalus, Benth. ...| Th. | | i+] | | 
iM. Stewartii, Wendl. nee.) aoe | | | f+ 
| Angianthus tomentosus, Wendl. =v, Seb +} {+ | 
| A. pusillus, Benth. ... fe Th. | +| 
=e brachyappus, Fey. MS. wo eee | t | | 
Gnephosis cyathoppapa, Benth. a ie is + voice 
B.|G. skirrophora, Benth... ; sed te | ea 
| Podotheca angustifolia, Cass. 2& itieOie | | |+i | | 
B.| Podolepis capillaris, (Steetz) Diels. ...| Th. | | j++ 4 
B. Pacichisysam apiculatum, 1). 3 ee iene | Choe eI | 
|H. ambiguum, Turcz. ae Sat Glee +| | | 
| H. Lawrencella, a. ve oy, Daven-| Th. | one 
PONTE TEE el a |e | fee (oat 
igs bracteatum, Andr.. ear Ch. | | ae) | | 
|H. semifertile, F. v. M. ... ote Eat | l+{ | f 
' Waitzia acuminata, Steetz ... eer fe a a | | foes 
| Helipterum polygalifolium, DC. pee e PaeT ae [| 
iH. floribundum, DC. aE > speedos i el a tt es 
| H. roseum, (Hook.) Benth., var. patens,| Th. | [ |+i+t | 
. Ewart : ees ear cos Ne| fe nome | 
|H. hyalospermum, F. v. M. | Bho Pe aep ye ees 
| H. strictum, Benth. ea | | |+j+i | 
| H. pygmeum, Benth. ope ag tice | 
| H. moscatum, Benth. se he, Peed wel 4 
|H. Humboldtianum, DC. Oe opr tay em 
| Cephalipterum Drummondii, Ae Gray ih eee Eee eg | 
peeucco Grevori. Wloivy: Moy to eh Thee poled] 
'S. brachyglossus, F. v. M. at is eee Spek ag ee 


DESCRIPTION OF PLATES. 


Prats XXXII. 

Fig. 1. General view on Nullarbor Plain showing typical 
bluebush (Kochia sedifoliaj. A donga is seen on the left. The 
trees in the donga are Acacia aneura, Pittosporum phillyraeoides, 
and Fusanus acuminatus. The soil of the plain shows white lime- 
stone fragments, also the flower heads of Cephalipterum Drum- 
mondit. 

Fig. 2. Transition region between plain and sandhill. The 
trees are not confined entirely to depressions (dongas), though 
one with a denser tree flora is seen to the left. The main tree is 


_ Acacia aneura, with occasional A. tetragonophylla and Eremophila 


Latrobei, etc. Atriplex vesicarium is the chief ground shrub, the 
soil here heing more sandy than on the plain (cf. fig. 1). Numerous 
annuals are present. 


564 


Pratt XXXII. 
General birdseye view over sandhill region looking west 
towards the Nullarbor Plain, which is seen on the horizon, taken 
from the top of a water tower 30 ft. above the crest of a sand 
ridge. In foreground sandhill mulga (A. linophylla) running to a 
flat with Heterodendron. Another low sand ridge is between this 
and the plain, where is mulga (A. aneura) and some Casuarina 
lepidophloia in dongas, @.g., extreme right. 


parte gg DY nS 


Prats XXXIV. 

Fig. 1. Typical sand ridge mallee vegetation. The mallees 
are EHucalyptus oleosa, E. leptophylla, and E. transcontinentalis. 
The tree in the centre is Myoporum platycarpum. Shrubs, Acacia 
ligulata, A. Randelliana, and Cassia eremophila. The ground at 
this season is almost bare. This probably represents the climax 
on sandhills. 

Fig. 2: View in ‘‘oak forest,’’ showing old trees of Casuarina 
lepidophloia with some natural regeneration. Bushes of Acacia 
Randelliana and Fusanus acuminatus. Ground at this time almost 
bare; the dead undershrubs appeared to be Kochia sedifolia. 


PusTteE XXXYV. 

Sandhill flora showing early stages in the succession in the 
foreground and looking across to mallee on the horizon. The trees 
in the foreground on either side are Grevillea stenobotrya. Below 
open sandhill succession with Leptospermum laevigatum, v. minus, 
Acacia ligulata, and Dodonaea viscosa, v. attenuata. The dis- 
tant vegetation is mallee of the sandhill] climax. 


PLuate XXXVI. 


Fig. 1. General view over salt lake to mallee-covered hills 
on horizon. The foreground shows open sandhill succession with 
Acacia ligulata and Fusanus acuminatus. The halophytic shrubs 
of the dry salt lake are seen round the margin and in the bed 
at the nearer and shallower end. Beyond the lake can be seen 
the halophytes, some passing into open sandhill flora, behind which 
is mallee. The branch of the tree in the immediate foreground, 
left side, is Grevillea stenobotrya, and illustrates the leaf habit. 
The tree is fruiting. 

Fig. 2. Salt lake near Soak showing dry bed with no vege- 
tation. Arthrocnemum, spp., form a fringe between the bed of 
the surrounding sandhills on which Acacia linophylla and Fusanus 
acuminatus can be seen. 


565 


ADDITIONS TO THE FLORA: OF SOUTH AUSTRALIA. 
No. 20. 


By J. M. Brack. 
[Read October 19, 1922. | 
Bi.  Prare XX XVII. 


GRAMINEAE. 


Stipa setacea, R. Br, var. latiglumis, n. var. Variat 
ligula 3-5 mm. longa, glumis vacuis latis, superiore sub-5-nervi, 
gluma florifera lata ad apicem angustata et breviter barbata, 
arista 25-35 mm. longa crassiuscula bis geniculata. 

Belair; Minnipa; Telowie Gorge. 


S. eremophila, Reader, var. dodrantaria, n. var. Variat 
gluma floriferaé angustiore, aristé 6-7’cm. longé usque ad 
dodrantem subplumosa. 

Birksgate Range, R. Helms. 


S. pubescens, R, Br., var. comosa, n. var. Variat gluma 
florifera circiter 4 mm. longa sericeo-villosa in comam albam 
aequilongam desinente. 

Marino; Jamestown; Melrose; Moolooloo. 


Eriachne ovata, Nees, var. pedicellata, n. var. Variat 
pedicellis capillaribus 5-8 mm. longis, gluma florifera superne 
villosissima sed non ciliata glumas vacuas paululum superante. 

Musgrave Range, S. A. White. 


CYPERACEAE. 


Cyperus exaltatus, Retz., var. minor, n. var. Variat 
umbella minore, spiculis 3 mm. longis 6-floris. 
River Murray. 


Scheonus tesquorum, n. sp. Perennis, caulibus fili- 
formibus compressis striatis 20-40 cm. altis, foliorum 
radicalium laminis filiformibus 6-18 cm. longis, bracteis 
caulinis: 2-3 distantibus, earum vaginis 5-20 mm. longis 


eylindricis fusco-rubris indiscissis, laminis filiformibus 1-4 cm. 


longis, spiculis fusco-rubris 6-7 mm. longis 2-floris lanceolatis 


| binis usque quaternis in fasciculos terminales et subterminales 
dispositis, setis hypogynis nullis; nuce obovoidea trigona 


~ alba. 


From between Mount Burr and Mount McIntyre to 


Nangwarry on the Victorian border. The types are four 


566 


specimens in the Tate Herbarium, without date or name of 
collector, but very probably aathered by Professor Tate him- 
self. One of them is marked for transmission to Baron von 
Mueller, but it appears, from enquiries at the National 
Herbarium of Victoria, that none was sent. The species 
stands nearest to S. apogon, Roem. et Schult. 


IRIDACEAE. 


*Moraea xerospatha, MacOwan, var. monophylla, 
n. var. WVariat folio radicali semper unico. 


‘In the typical South African form of this little plant, 
very common in our southern districts, and especially near 
Adelaide, the number of leaves is given by Baker as 3-4, and 
by the Kew authorities as 1-4. 


CHENOPODIACEAE. 


Chenopodium carinatum, R. Br., var. melanocarpum, 
n. var. Variat perianthio fructifero demum nigrescente, 
ejus segmentis pilosioribus margine contiguis semen tegentibus 
subacute carinatis. Accedit Ch. cristato. 


Flinders Range; Far North and North-West. Also in 
Western Australia and at Broken Hill, New South Wales. 


Chenopodium microphyllum, F. v. M., var. desert- 
orum, no. var. Variat caulibus erectioribus et crassioribus, 
foliis ovatis vel rhomboideis crassis supra concaviusculis infra 
dense farinosis 5-12 mm. longis, spicis densius vestitis folia 
superantibus (10-15 mm. longis), staminibus 5. 

Murray lands; Port Augusta westward to Ooldea. 

This variety, with thick, often rhomboid, almost papillose 
leaves and longer spikes, looks very distinct, but some speci- 
mens from Baroota, between the Flinders Range and Spencer 
Gulf, are in some respects almost intermediate between the 
type and the variety. 


Bassia ventricosa, n. sp. Fruticulus ramosus, ramulis 
albo-tomentosis, foliis lineari-clavatis sessilibus acutis 
sericeis demum glabrescentibus 5-15 mm. longis, floribus 
solitariis, perianthii fructiferi tubo parce tomentoso subgloboso — 
3 mm. diametro ad basin oblongam vix excavato, limbo lanato 
sat longo, spinis 4 (rarissime 5), quarum duabus 3-5 mm. longis 
ceteris valde brevioribus una nonnunquam adnata vel paene 
obsoleta omnibus rigidis subdivergentibus plus minusve 


pilosis, pericarpio superne indurato, semine oblique | 


horizontali. 


TT Ee ee 


567 


Port Augusta and Lake Torrens to the Far North; also 
in the western part of New South Wales. This widely dis- 
tributed species is easily distinguishable from its allies by its 
almost. globular hairy perianth-tube and its four short un- 
equal slightly divergent spines. 


Bassia limbata, n. sp. Fruticulus ramosus tomento 
denso albo-cinereo paene lanato tectus, foliis lineari-clavatis 
sessilibus 10-20 mm. longis, floribus solitariis, perianthii 
fructiferi tubo subcylindrico 3 mm. longo ad apicem 4 mm. 
lato ad basin oblongam vix excavato cum limbo erecto 
aequilongo aibo-tomentoso, spinis 2 divergentibus rigidis 
erassiusculis 8-12 mm. longis usque supra medium tomentosis, 
tertia spina vel tuberculo minuta, semine horizontali. 

Leigh Creek, Parachilna, Mount Parry (Flinders Range) ; 
also near Broken Hill, New South Wales. Allied to B&B. 
bicornis, (Lindl.) F. v. M., but the latter has a much larger, 
harder, and more woolly ‘perianth-tube, fiercer spines, less 
conspicuous limb, and has no third spine or tubercle on the 
inner face. The basal area of attachment, oblong and 


-searcely hollowed, in fact only a broad groove, is much the 


same in both species. 


Bassia decurrens, n. sp. Fruticulus suberectus, ramis 
lanatis denique glabrescentibus, foliis lineari-subteretibus 
acutis sessilibus parce pilosis 10-15 mm. longis, floribus 
solitariis, perianthii fructiferi tubo subcompresso glabro vel 
circum basin lanulato laevi costato-sulcato circ. 3 mm. longo 
latoque ad basin ovatam paulo excavato, spinis 2 glabris 
divergentibus basin versus dilatatis 6-8 mm. longis, quarum 
una in 3 spinas brevissimas vel tubercula decurrit, limbo 
tubum aequante ad apicem truncato et ciliato, semine 
vertical. 

Near Port Augusta; also in western New South Wales. 


Differs from the other species with vertical seeds in its smooth 


ribbed glabrous perianth-tube, with two broad-based divergent 
spines of which one terminates at the base in three very short 
spines or tubercles, while the base of the tube is ovate or 
almost orbicular and only slightly oblique and hollowed on 
the inner face. 


Bassia paradoxa, (R. Br.) F. v. M., var. latifolia, 
n. var. Variat folus 15-25 mm. longis 5-8 mm. latis dense 


| tomentosis, capitulis 15 mm. diametro, spinis (in paucis 
_ Speciminibus quae adsunt) ad 5 cornua brevia obtusa reductis. 


Strzelecki Creek, S. A. White. The five short obtuse and 


rather soft horns are very different from the usually sharp 
_ Nigid spines of typical B. paradoxa, but it is certain that con- 
siderable variation exists in the length and texture of the 


568 


dorsal appendages even in specimens which are otherwise 
typical. The new variety can be at once distinguished by its 
very broad thick and soft leaves. My specimens are too few 
to ensure certainty as to the appendages being ona of 
the form described. 

In dealing with the Bassias, I have had the advantage 
of consultation with Mr. J. H. Maiden, Government Botanist, 
and Mr. R. H. Anderson, botanical assistant at the National 


Herbarium, Sydney. Mr. Anderson is engaged on a much- — 


needed revision of the Australian Rassias. 


Atriplex leptocarpum, F. v. M., var. acuminatum, 
n. var. Folis obovatis plerisque sinuato- dentatis; bracteolis 
fructiferis 5-8,mm. longis, lobis (partibus liberis) ‘acuminatis 
tubo indurato fere aequilongis nonnunquam in utroque 
margine minute unidentato et saepe ad basin 2 parvis tuber- 
culis dorsalibus instructis. 


Tarcoola; Ooldea. 


Babbagia acroptera, F. v. M., et Tate, var. deminuta, 
n. var. Variat grandiore ala crassa rubella oblongo-incurva 
vix 2 mm. longa, altera minima vel fere obsoleta. 

West of Port Augusta. 


Kochia scleroptera, n. sp. Fruticulus subereetus, ramis 
albo-tomentosis, foliis linearibus acutis sericeo-villosis 6-12 
mm. longis nervo medio saepe conspicuo infra, floralibus saepe 
caducis, floribus in longas spicas confertis, perianthio 
fructifero. valde depresso sub tomento dense lanato obtecto 
4-5 mm. diametro 5 alis brevibus obtusis rigidis crassiusculis 
horizontalibus comprehensis, tubo brevissime convexo vix 
1 mm. longo 2 mm. lato infra alas ipsas. 

Arkaringa and Alberga Creeks. This species is ‘only 
known by two specimens collected by R. Helms on the Elder 
Expedition in 1891, and one obtained by Miss Staer in 1913 
at Todmorden Station, on the Alberga. Like XK. brevifolia 
it has five distinct equal horizontal wings without appendages, 
but differs entirely in the thickness and rigidity of the wings 
and in their dense woolly clothing. The perianth bears some 
resemblance to that of A. lanosa or of Bassia sclerolaenoides. 


UMBELLIFERAE. 


Uldinia, n. gen. Floribus paucis breviter pedicellatis 
in umbellam simplicem pedunculatam conjunctis, calycis 
dentibus obsoletis, petalis ovatis obtusis leviter imbricatis, 
stylis brevibus, fructu ovato a latere valde compresso basi 
emarginato mox bipartibili, mericarpiis 5-jugis, jugo dorsali 
aculeis uncinatis divaricatis biserialibus marginato, jugis 


> 


dud 
ris 


569 


intermediis prominulis medianis parcius aculeatis, utroque 
jugo intermedio in alam divaricatam lanceolatam uncinato- 
ciliolatam 3-5 mm. longam ad apicem desinente (alis eas 
pedium Mercurii simulantibus), jugis marginalibus prominulis, 
vittis nullis, carpophoro obsolescente setaceo uno mericarpio 
adnato et cum eo deciduo, foliis palmatipartitis petiolatis, 
petiolo basin versus subdilatato pilis longis ciliato ad basin 
ipsam fimbriato sed non rite stipulato. 


Uldinia mercurialis n. sp. (Tab. xxxvii.). Herba 
annua, caulibus prostratis rigidis plerisque simplicibus glabris, 
foliis radicalibus longe petiolatis, lamina ambitu  sub- 
orbiculari-cordata 10-15 mm. longa parcissime pilosa tri- 
partita, segmentis ovato-cuneatis obtuse trifidis vel lobato- 
incisis, petiolo 10-30 mm. longo, foliis caulinis valde minoribus 
plerisque oppositis, umbellis 4-floris, pedunculis robustis 
5-7 mm. longis axillaribus, petalis caeruleis 1 mm. longis, 
involucri bracteis 4 lanceolatis ciliatis circiter 3 mm. longis, 
mericarpiis minute papillosis 4 mm. longis vix 2 mm. latis 
minus quam | mm. crassis, alis divaricatis 2-4 mm. longis. 


This curious little plant, collected by Mr. E. H. Ising 
along the railway line at Ooldea in September, 1920, does not 
seem to fit into any of the existing genera of the tribe 
Hydrocotyleae. Tt has the habit of Didiseus, but differs in 
the absence of a free persistent carpophore. In the somewhat 
dilated and ciliate base of the petiole it resembles D. glauci- 


_folius. The carpophore of Uldinia is setaceous and adnate to 


the slightly grooved face of the narrow commissure of one of 
the two mericarps. It is fragile at base and falls off with the 
mericarp to which it is attached. From Hydrocotyle it differs 
in the deeply-cut leaves, the absence of stipules, the imbricate 
petals, and the dry station; from Trachymene, DC., in the 
absence of a free persistent carpophore, in the petals not 
inflexed, in the simple umbels and the dilated base of the 
petiole; from Cente/la in the lesser number of ribs on the 
mericarp and the divided leaves. As regards the two new 
Australian genera created by Domin in 1908 (in Beihefte 
zum Botan. Centralblatt, xxiu., Abt. i1., 291-4), it differs 
from NVeosciadium in the flattened fruit, deeply-cut leaves, 
and in the absence of free stipules. (N. glochidiatum, 
Dom. = Hydrocotyle glochidiata, Benth. ; Centella glochidiata, 


_ Drude.) From the other new genus, Homalosciadium, it 
differs in the adnate deciduous carpophore, and from both 
_ of these genera in the few-flowered umbels. (H. verticillatum, 


Dom.= Hydrocotyle verticillata, Turez.; Centella homalo- 


_carpa, Drude.) The hooked prickles or bristles which stand 
oR 


~ 
: 
‘he yl 
—— 


570 


in two rows along the narrow keel or dorsal rib of the meri- 
carp and are scattered along the intermediate ribs, differ 
markedly from those of Neosciadiwm, where they consist of 
straight slender bristles with several ‘short reflexed barbs or 
hairs near the summit, whereas in U/dima they are simple, 
stout, and hooked at the end. Still more remarkable are the 
two lanceolate wings attached to the summit of each inter- 
mediate rib and spreading outwards at right angles to the 
flattened sides of the mericarp. By their shape and position 
they recall the wings with which classic legend adorned the 
feet of Mercury. As far as my knowledge goes, they do not 
occur in any other umbelliferous plant. They also have small 
hooked prickles along the margin, so that the fruit appears — 
well adapted for transport either by animals or by the wind. 
The divaricate wings and the hooked prickles should perhaps — 
be ranked rather as specific than generic characters, but even — 
in that case the other peculiarities of the plant appear suffi- 
cient to necessitate the creation of a new, although probably 
monotypic genus. 


The name of the new genus is derived from ‘“‘aildilnga 
gabi,’’ the native name of ‘‘Ooldea Water,’’ more generally 
known as the Ooldea Soak, and about three miles from the 
Ooldea Railway Station. 


MYOPORACEAE. 


Eremophila pentaptera, n. sp. (Tab. xxxvi.) Frutex 
humillimus glaber circ. 30 cm. altus, caulibus erectis, foliis 
alternis crassis subplanis oblongo-cuneatis sessilibus ob- — 
tusissimus 10-35 mm. longis 4-8 mm. latis, floribus solitariis 
subsessilibus, pedunculis brevissimis erectis obconicis acute 
quinquangulis circ. 5 mm. longis, calycis segmentis aequalibus 
circ. 12 mm. longis glabris lanceolato-acuminatus sed. obtusis 
valde imbricatis secus dorsum acute carinatis vel anguste 
alatis in pedunculum brevem decurrentibus, corolla violacea 
25-35 mm. longa exterius glabra in faucibus alba lanata 
maculis fulvis obsité, tubo ad basin cylindrico sursum sensim 
dilatato, omnibus lobis rotundatis et tubo aequilongis (exceptis 
2 supremis brevioribus) infimo truncato 14-18 mm. lato, 
staminibus inclusis, ovario conico glabro, stylo pilosulo, ovulis 
2 in utroque loculo, fructu non viso. 

This lowly Eremophila was discovered by Professor F. 
Wood Jones in September, 1922, on flats near Miller Creek, 
about 60 miles (100 km.) north-east of Kingoonya Railway 
Station. It appears to be local in its distribution. The ovary 
has two cells, each with two ovules, and the corolla-tube has 
a cylindrical base, in which respect it agrees chiefly with 


ee 


~ 


im 
q 
oe 
x 


571 


Pholidia, but the size and shape of the upper part of the 
corolla and the large thick leaves belong rather to Hremo- 
phila. In any case, the desirability of uniting the two genera 
seems now to be generally conceded. In the broad, fleshy, 
rather large, and very obtuse leaves, and the sharply-keeled. 


or narrowly-winged calyx-segments, which run down into an 


acutely 5-angled peduncle so short that it appeared to be 
merely the contracted base of the calyx, the new species is 
well distinguished. 


DESCRIPTION OF PLATE XXXVII. 
1. Eremophila pentaptera, n. sp. A, branch. 8B, corolla 


; spread open. C, dangled peduncle and pistil, D, vertical section 


of ovary. HE, calyx and peduncle. 


2. Uldinia mercurialis, n. sp. F, radical leaf. G, upper part 
of stem. H. mericarp. I, transverse section of fruit. 


572 


TYPES OF SPECIES OF AUSTRALASIAN POLYPLACOPHORA 
DESCRIBED BY DE BLAINVILLE, LAMARCK, DE ROCH- 
BRUNE, AND OTHERS, NOW IN THE MUSEUM D'HISTOIRE 
NATURELLE, IN PARIS. 


By Epwin Asupy, F.L.S., M.B.O.U. 
[Read October 19, 1922.] 


The following is a reswmé of the results of an examina- 
tion recently made by the writer of the collections of Aus- 
tralasian Polyplacophora under the care of the Laboratoire de 
Malacologie Rue de Buffon, Paris. 

The writer’s warmest thanks are due to Professor Joubin 
for permission to examine the collections, and to Dr. Ed. 
Lamy, for not only placing the extensive collections at his 
disposal, but also for much help in the identification of the 
specimens from which Blainville and other writers made 
their original descriptions. 

In offering the within notes on these collections, the 
writer is conscious of limitations due to the shortness of the 
time at his disposal entirely precluding the possibility of 
checking through his rough notes before transcribing them. 

The fortunate rediscovery of some of the lost types, 
notably of Blainville and Lamarck, on which so much has 
been written by Dr. Pilsbry, Mr. Tom Iredale, and others, 
will, I feel sure, be appreciated by all workers. 

Fairly full notes have been given of a good deal of 
material of less importance than the types before referred to. 
This has been done because the writer had an unique oppor- 
tunity of comparing the specimens with those of his own 
collection which he brought to Europe for this purpose, and 
which is undoubtedly the most complete collection of Aus- 
tralian chitons that has up to the present been made. The 
references given are not complete, but sufficiently so for the 
purposes of this paper. As far as possible the notes have been 
arranged in the order of modern classification. 

Callochiton dentatus, Spengl., Australe. One specimen 
on card. On the back is marked ‘‘fulgetrum, Reeve.” It is 
very worn, but I have no doubt it is C. platessa, Gould. 

Lepidopleurus fodiatus, Rochebr. Type (Bull. Soc. 
Philom., 1880-81, p. 120). The card on which these shells 
are mounted is marked ‘‘/s. (Radsella) fodiatus, Rochebr.”’ 
Also, there are several separate valves in spirit marked on 
label ‘‘Zs. tigrinus, Kraus.’’ Other notes on the label, ‘“New 
Holland, M. Verreaux, 1842. Type, M® 108.”’ 


_ 


42 - 


{4 
iy 


573 


This shell has very large scales grooved with very fine 
striae. Lateral area 11 radial ribs, median areas covered 
with flattened, wavy ribs which are so extremely bridged as 
to approximate to the sculpture of a Callistochiton. 


I have never seen this shell in Australia and am con- 
fident the locality given is incorrect. 


Stenochiton (Chiton) longicymba, Blainville. Type (Dict. 
Sc. Nat., xxxvi., 1825); Stenochiton juloides, Ad. and Ang. ; 
Schizochiton nympha, Rochebr., non Chitow longicymba, B1., 
of Quoy et Gaimard. : 

The full particulars of the steps that led to the identi- © 
fication of Rochebrune’s type of Schizochiton mympha with 
Blainville’s lost type of C..longicymba are fully given in a 
paper by the writer which is being published by the Malaco- 
logical Society, London. 


Ischnochiton (Chiton) hneolatus, Blainville. Type (Dict. 
Se. Nat., vol. xxxvi., p. 541, 1825). See ‘Review of Chiton 
e<rispus, Reeve, by Ashby (Trans. Roy. Soc. S. Austr., vol. 
xliv., 1920, pp. 272-275); non feneolatus, Blain., of Iredale 
and May; haddoni, of Pilsbry; 7. erispus, Reeve, of Bednall, 
Torr. In my paper (/.c.) I recognized Blainville’s Chiton 
dineolatus in the shell we had known as /. crispus, Reeve, 
common to South Australia, Victoria, and Tasmania; and 
later, I received from Major Paul Dupuis, of Brussels, co- 
types of Blainville’s lineolatus, collected by Peron et Lesueur 
ain King Island, one of the specimens being marked inside 


“Tle King.” This confirmed my previous conclusions. Of 


the two specimens mounted on the card marked ‘‘Type 
Peron et Lesueur, Ile King,” the larger shell is smooth and 
shiny, has no radial ribbing, but shows near the insertion 
plates, broken, shallow, concentric ribbing. The dorsal area 
is almost smooth and polished, the fine decussation, although 
present, is inconspicuous. The lateral and median areas are 
much less coarse in sculpture than is the case with /schno- 


_chiton tredaler, Dupuis, which is the shell that Iredale had 


concluded was Blainville’s lineolatus; the jugum in the type 
is not so rounded as in the latter species, but shows a fairly 
sharp angle and also a single flame mark bordering the dorsal 
area. Except the flame mark the type corresponds with the 
shell given me by Major Dupuis before referred to. In the 
second and smaller specimen on the card, the radial ribbing 


1s present on the anterior and posterior valves and in the 


lateral areas of the median valves; altogether the sculpture 
is much stronger than is the case with the larger specimen. 


_ While this sculpture approaches /. iredalei, Dupuis, the 


marked jugal ridge separates 1t from that species, and the 


574 


sculpture of this area is less strong than on that species. The 
girdle, while showiag stains, is certainly a pale girdle. 

There are on three other cards marked ‘“‘Peron et 
Lesueur, [le King,’ one, two, and three specimens, respec- 
tively. In all the sculpture is coarser than is the case with 
the larger one on the card marked ‘‘type.’’ While it is pos- 
sible that some of them may be juvenile specimens of 
Heterozona sub-viridis, Ire. and May, I could not distinguish 
any pointed large scales that are so characteristic of that 
species. 
In conclusion:—This investigation determines ute facts 
that (1) Iredale was wrong in identifying the shell we used 
to know as. [schnochiton contractus, Rv., and now known 
as I. zredalei, Dup., with Blainville’s Chiton lineolatus, 
(2) The larger specimen on the type card, which I accept as 
the type, is undoubtedly the shell we used to know as J. 
crispus, Rv. (3) While it is possible that some of the specr- 
mens brought from King Island by Peron and Lesueur, and 
mounted on the separate cards may be juvenile forms of 
H. sub-viridis, Ive. and May, their present condition makes 
accurate determination difficult. 

Ischnochitow (Lepidopleurus) campbell, Filhol. Type 
(Comptes Rendus, xci., p. 1095, 1880). Iredale in Traas. 
N. Z’d. Inst., vol. xlvu., 1914, p. 419. The type is fromm 
Campbell Island and is a half-grown specimen similar to one 
of the same size in my own collection. 

Ischnochiton melanterus, Rochebrune. Type (Bull. Soc. 
Philom., Paris, 1883-84, p. 137), from Campbell Island. 
This is conspecific with the preceding species. 

Ischnochiton (Chiton) tessellatus, Quoy et Gaimard. 
Type (Voy. de l’Astrol. Zool., iii., p. 396; Atlas, t. 7am 
f. 43-47). This specimen is mounted on a card and marked 
“Tl des Kangaroo,’’ but in Pilsbry the locality is given 
Port Carteret, New Ireland. On the back of the card there 
is the note, “‘“C. cymbium, Dufrizai. M.S.S.’’ This shell has 
girdle scales that are large, bead-like, polished, and almost 
smooth at apex; elsewhere very finely striated. Anterior and — 
posterior valves and lateral areas of median valves show 
strong, slightly broken, radial ribs 5 or 6 in number. The 
median areas closely packed with narrow, well-defined, wavy, 
longitudinal ribs, curiously pectinated. I have never seem 
this shell in Australia. 

Ischnochiton er sulcatus, Quoy et Gaimard. Type 
(Voy. de 1’ Astrol., 111., p. 385, t. 75, f. 31- -36). The card is 
marked “Codie urvillei, Rochebrune’”’ ; also ““M. My 
Quoy et Gaimard. Type, from Port du Roi George.”’ It 
is a strongly-marked specimen of the shell we now know as 


; 


575 


I. contractus, Rv. The synonymy is given by Iredale and 


| 
| 


May (Proc. Mal. Soc., vol. xii., pts. 1. and ii., Nov., 1916) 
as follows:—J. contractus, Rv., 1847; C. sulcatus, Quoy et 
Gaim., 1834, non Wood, 1815; C. decussatus, Rv., 1847; 
C. castus, Rv., 1847; Lepidopleurus speciosus, Ad. and Ang., 
1864; Gymnoplar urvillet, Rochebr., 1881. 
_ Plaxiphora (Chiton) biramosa, Quoy et Gaim., 1833. 
Type, New Zealand (Voy. de 1’Astrol., ill., p. 378, pl. 74, figs. 
12-16). This specimen is very badly eroded, showing no 
sculpture, only a few zebra-like markings on a brown ground. 
Plaxiphora (Chitow albidus) albida, Blainville. Type, 


le King (Dict. Sc. Nat., vol. xxxvi., p. 547, 1825). There 


are six valves in tube marked type, M, 36a, anterior, pos- 
terior, and four median valves. These were disarticulated 
by Dr. J. Thiele, and, as stated by him, are very bleached and 
eroded, but one of the valves shows distinct wrinkling, and 
all show, near margins, growth-lines and the usual, although 
faded, green and brown dashes or bands. 

Plaxiphora (Chiton costatus) costata, Blainville. Type, 
Port du Roi George (Dict. Sc. Nat., vol. xxxvi., p. 548). The 
“specimens, which are dissected, are also labelled “‘Chacto- 
pleura costata, Bl.’’ This is the usual ribbed and wrinkled 
form of Plax:phora found in South Australia. The anterior 
diagonal rib is well defined and a large portion of it smooth; 
the posterior margin of the median valves is not well defined. 
‘The wrinkling is very marked in zigzags. The usual green 
colour markings are present. 

Plamiphora (Chiton glaucus) ek Quoy et Gaim. 
Type, Van Diemen (Voy. de 1’Astrol. Zool., ii1., p. 376, 
t. 74, f. 7-11, 1834); marked in (Quoy or Gaimard’s) hand- 
writing as from Van Diemen. There are only four valves, 
which belong to the smooth form, without wrinkles. J have 
similar specimens from Tasmania. The shell is~a good deal 
eroded, but there is sufficient ie show that, when perfect, it 
was unwrinkled. 

Comment.—Dr. Thiele ani correctly recognized de 
Blainville’s C. costatus in the wrinkled form of Plaxi phora, 
Tanging from Victoria to Western Australia, which appears 
to have been described by Sowerby under the name of P. 
petholata. But he concluded that the bleached and eroded 
Specimen described by Blainville under the name of (. alhidus 
must be the smooth form described by Quoy et Gaim. under 
the name of (. glaucys. The discovery of defined wrinkling 
on one of the valves of Blainville’s type of C. albidus dis. 
proves this. I am satisfied that his albidus and costatus are 

conspecific, the former being a worn and bleached specimen of 
the latter. As albidus has page priority, it must stand as 


576 


the name of the wrinkled shell, and Piaxiphora costata, Bl., 
as a synonym thereof. Whether the whole of the forms of 
Plaxiphora found in Southern and Western Australia are all 
referable to one very -variable species or not, must be left to. 
future investigation. For the present I purpose to include all 
wrinkled forms from Victoria and Tasmania aud westward 
under the name of P. albida, Bl., with P. conspersa, Ad. and 
Ang., as a subspecific name for the extremely sculptured form 
which has its headquarters at Port Lincoln. The unwrinkled 
forms I would refer to P. glawea, Quoy et Gaim., of which the 
type is from Tasmania. 


Plaxiphora varipilosa, Blainville. Type (no locality). — 
This is a disarticulated specimen showing no sculpture and 
is very smooth and polished. More rounded (except near the 
beak, which is raised) than is the case with Australian repre- 
sentatives of this genus. It is decorated with longitudinal 
brown streaks, but has none of the typical markings that are 
so characteristic of Australian specimens; it is evidently not 
an Australian shell. 


Cryptoplux montanoi, Rochebrune. Type, in spirit, Is. 
Soulon, Drs. Montano et Ray (Pilsbry gives locality as 
Leucon, Borneo), No. 74, 1880 (Bull. Soc. Philom., Paris,. 
1881-82, p. 1901), is marked “C’. oculatus, Q. et G.” This 
specimen is well preserved in spirit and is conspecific with a 
specimen in my own collection which is also marked “OC. 
oculatus, Q. et G.,” and is from I. Sula. Both these differ 
from specimens marked “C. larvaeformis, (Blain.) Burrow, 
1815,’’ in that the first three valves are circular and not oval,. 
as is the case in the latter. In all other respects they seem 
alike. We were not able to find the type of C. oculatus, 
Quoy et Gaim., so cannot. say whether these two are con- 
specific. 


Cryptoplax (cryptoconchus) larvaeformis, (Blain.) Bur- 
row (Elem. of Conch., pp. 190, 191, t. 28, f. 2, 3, 4, 19TaR 
I saw a card with old label, ‘‘I. O. Lisse. Ch. laevis, Lam@ 
Habite les Mers de la Novelle-Holland,’’ determined later 
as C’. larvaeformis, Burrow. This specimen is similar to others 
in the same collection from New Caledonia and Tonga Tabu. 
I have noted that all these resemble my shell and Rochebrune’s 
montanoi, except that in them valves 1 and 2 only are round 
oval, whereas in the latter the three first valves are almost 
circular. The girdle spicules of all are very similar. | 


Cryptoplax lamarcki, Rochebrune, from New Caledonia, 
marked ‘‘co-type.”’ This specimen appears to correspond with 
C. larvaeformis, Burrow, in that the anterior valve is oval 
and not circular, as in mondawoi, Rochebr. 


v 


q 


577 


.Cryptoplax torresianus, Rochebrune. Type (Bull. Soc. 
Philom., Paris, 1881-82, p. 195, 1881). The following are 
copied from Rochebrune’s MS. notes :—‘‘Chitonellus striatus 
{Rv., C. Icon., pl. 1, sp. 4), long. O. 060, lat. O. 004, M.M., 
non C. striatus, Lam. Hab., detroit de Torres. Mus., Paris.’’ 
I compared this type with my shell from Sydney and found 
them conspecific; my largest specimen is a counterpart of the 
type, which is numbered N. 13, 8. 

Cryptoplax (Chitonellus) laevis, Lamarck. Type (de 
Lamarck animaux sans vertebres, vol. 7, Mollusques). Lab. 
de Malacologie, K. 82. Oscabrelle lisse, De Blainville, 
Malac., pl. 87, f. 5. Hab. les Mers de la Novelle-Hollandiae. 
Peron et Lesueur. I saw type marked as such in Roche- 
brune’s handwriting. The same specimen is marked “‘type of 
Cryptoplax (Chitoneilus) lamarchi, Rochebr.,’’ and the card is 
marked ‘‘Peron et Lesueur, N. 1° 3,’ and agrees with figure, 
pl. 87, fig. 5, Blain., Manu. de Malacologie. This type shell 
measures 49x12 mm., and, as just stated, corresponds with 
the figure. Nearly the whole of the shell is eroded and the 
girdle is denuded of spicules, except on that portion com- 
mencing opposite valve 7, where the spicules are fortunately 
still in evidence. These spicules are very peculiar, being 
very widely spaced, short, blunt and rounded, quite distinct 
from any species I have seen from either the south or the 
east coast of Australia; neither does it agree with the speci- 
men of Cryptoplax I collected at Rottnest, in Western Aus- 
tralia. On the other hand, the small specimen I collected 


at Yallingup, in that State, in October, 1920, and provision- 


ally referred to as C. hartmeyer:, Thiele—see my paper (Trans. 
Roy. Soc. S. Austr., vol. xlv., 1921)—may be a juvenile of 
this species. Should this identification ultimately prove cor- 
rect, UC. hartmeyeri, Thiele (the type of which I have never 


seen), will probably be proved to be conspecific, and name will 


be a synonym of C. laevis, Lam. 

Cryptoplax (Chitonellus) striatus, Lamarck. Type (An. 
S. Vert., vi., p. 317, 1819): The type specimen is marked 
in Lamarck’s handwriting, ‘‘Oscabrelle striée, Chitonellus 
striatus. Tle aux Kangaroo.’’ The type measures 46 x10 


mm.; valves 5, 6, and 7 would be slightly spaced if the speci- 


men had been carefully dried. The sculpture is similar to 
the common South Australian shell, and although very few 


spicules are left on the girdle, those that remain correspond 


with the South Australian species, which is quite natural, as 
Kangaroo Island, in that State, is the type locality. 


Comments.—The rediscovery of Lamarck’s two lost types 


_ is due to the very careful search made by Dr. Lamy. I was 
told on arrival that these types were not in the Museum in 


578 


Paris, and were probably in Gemeva. But on calling attention 
to the reference in the original description to their deposition 
in Paris, Dr. Lamy turned up Rochebrune’s MS. and found 
that when those notes were written Lamarck’s types were still 
in Paris. A further search was at last rewarded with their 
recovery. Pilsbry’s figures in pls. 9 and 11, Man. Con., vol. 
xv., are, I have no doubt, drawn from New South Wales 
specimens. I agree with Mr. Tom Iredale that these northern 
shells are a distinct species, and not Lamarck’s striatus. As 
a result of the foregoing investigation we are able to 
recognize : — 

(1) Cryptoplax laevis, Lamarck=C. lamarcki, Rochebr ; 
probably =C’. hartmeyers, Thiele, and is only known for Wes- 
tern Australia. | 

(2) Cryptoplax striatus, Lamarck, found in Victoria,. 
Tasmania, South Australia, and Western Australia. 

(3) Cryptoplax torresianus, Rochebrune. . Found from 
Port Jackson northwards to Torres Straits. 

(4) Cryptoplax guna, Reeve. Occurring in South Aus- 
tralia and having probably a range of habitat coextensive with 
C’. striatus, Lam., from which species it is easily distinguished 
by its dense, very short, and slender spicules. 

Three further species are all said to occur in Australian 
waters :—C. oculatus, Quoy et Gaim.; C. burrow, Smith; 
C. michaelsem, Thiele. The two first in the tropical waters 
of Queensland, and the latter in Shark Bay, Western 
Australia. 

Acanthochiton sueurn, Blanville.. Type (Dict. Se. Nat., 
Xxxvl., p. 553, Blainville). There are two specimens with: 
original label ‘‘Port Roi George.’? They are undoubtedly the 
shell we have known as A canthochiton asbhestoides, Smith. The 
better of the two is similar to a pale specimen I collected at 
Port Lincoln, in South Australia. 

Acanthochiton. jacundus, Rochebrune. Type (Bull. Soc. 
Philom., 1881-82, p. 194). There are a number of specimens 
in spirit which are conspecific with preceding species, all much 
worn. 

Acanthochiton. violaceas, Quoy et Gaim. Type (Voy. de 
lV’ Astrol., iii., p. 403, t. 73, f. 15-20), New Zealand. These 
are similar to specimens in my own collection from Doubtless’ 
Bay, New Zealand. 

Acanthochiton violaceus, var. papillo. Type. On another 
card marked “‘Quoy et Gaimard, 1883, N.Z.,’’ is a dissected 
specimen with anterior valve missing. All valves smooth and 
of peculiar shape. I am rather doubtful whether this is refer- 
able to the same species. It is referred to in Voy. de 
l’Astrolabe at top of page 520 under the name papillo. 


575 


Cryptoconchus (Acanthochites) meonticularis, Quoy et 
“Gaim. Type from New Zealand (Voy. de ]’Astrol., p. 106, 
+. 73, f. 30-35, 1834). This is undoubtedly conspecific with 
_CUryptoconchus porosus, Burrow (Elem. Conch., p. 189, t. 28, 
mi 1, 1815). 

Cryptoconchus stewartianus, Rochebrune. Type (Bull. 
Soc. Philom., Paris, p. 194, 1881-82). Type is in spirit and 
is evidently conspecific with preceding species. 
Acanthochiton zelandicus, Quoy et Gaimard. Type, in 
"spirit (Zool. Voy. de l’Astrol., i1., p. 400, t. 73, f. 5-8, 1834), 
_ marked on label “‘M. Quoy et Gaimard, 1833.” The shell is 
decorated with flat, rounded pustules. The dorsal area is 
partly smooth, but the rest of the area shows longitudinal 
“striae. They are similar to shells in my own collection from 
Auckland Harbour. 

Acanthochiton (Loboplax) stewartiana, Thiele. Type is 
in spirit and marked ‘‘Collected by Filhol, He Stewart.” The 
following are my notes:—Anterior valve ‘decorated with five 
well-raised rays formed of largish, elongate, flat pustules. 
_ Median valves with a diagonal fold and decorated with rows 
of diagonally-placed, raised, oval, flat pustules; much like 
A. granostriatus, Pilsbry. "The posterior valve is very dis- 
“tinct, dorsal area pinnatifid, in the front of mucro pustules 
are similar in character to those of the median valves. Mucro 
‘raised, posterior and distinct. Area behind mucro flat to 
concave. A small specimen in my collection, from Wel- 
lington, is similar in sculpture but has not the strong rays 
an the anterior valve. 

Note.—Several other Acanthochitons in the collection in 
t aris will be dealt with later. 

ie Rhyssoplax (Chiton) canaliculatus, Quoy et Gaimard. 
lmetype (Voy. de 1|’Astrol. Zool., ni., p. 394; Atlas, t. 75, 
 f. 37-42, 1834), marked ‘‘Voy. Astrol., 1829, New Zealand.” 
ere are no scales left on girdle of type. The sculpture is 
‘similar to specimens so named in my collection. 


Sypharochiton (Chiton) pellrs- -serpentis, Quoy et Gaimard. 
type, in spirit (Voy. de l’Astrol. Zool. in., Moll., p. 381, 
. 741, f. 17-22, 1834); label in handwriting of de Blainville 
reads, “Oscabricon a Serpent, pl. 741, fig. 17-22, New Zealand, 
Bictrclobe.’ There are three specimens of this well-known 
shell. In all there is very little sculpture in the median areas, 
_but the lateral areas are quite normal. One of the three is 
quite as carinated as S. sinclairi, Gray. 


Inolophura (Chiton) hirtosus, (Peron MS.) Blainville. 


| Dype (Dict. Sci. Nat., xxxvi., p. ‘546, 1826), Dr. J. Thiele 
. (! auna S.W. Austr. ie 399, 1911), Dupuis (Ex. Bull. Mus. 


580 


Hist. Nat., 1917, No. 7, p..1, 2), and (.c., p. 7, 19182) eam 
latter point out that in the Paris Museum are two specimens 
collected by Peron; the one marked type, which I call (a), 
is conspecific with Lzolophura (Chiton) georgiana, Quoy et 
Gaim. The other, which I call (6), is marked co-type, was 
identified by Dr. Thiele as conspecific with Acanthopleura 
spumger, Sow. The following are my notes on the type speci- 
men, which is disarticulated. The card is marked ‘‘Chitow 


hirtosus, Peron=Chiton georgianus, Quoy et Gaimard, [le 


King”’; and has labels at the back which read, ‘“‘See pg. 533, 
Bull. du Mus. d’Histoire Nat. (in Blainville’s handwriting), 
hirtosus, Bl.’’; in Peron’s handwriting, “‘hirtosus, Ile King”’ ; 
and in Quoy or Gaimard’s handwriting, ‘‘Aculsatum.”’ 

The large median valve measures laterally 30 mm. The 
sculpture is almost entirely eroded, broken and beaded growth- 
lines are slightly visible in the lateral areas, and what little 
sculpture remains on the rest of the valve consists of concentric 
ridges. The anterior valve is slit and propped, but the insertion 
plate is absent. or modified in the manner characteristic of Z~ 
georgiana, Quoy et Gaim.; the girdle scales are also similar 
to that species. There is a note on the back of the card, 
“Dr. Thiele det. this is undoubtedly Quoy et Gaimard’s 
shell.’ In this determination I fully concur. I have no 
doubt the specimen came from Port du Roi George, and not 
from Ile King, as marked on card; but until a careful search 
for Chitons be made on that island, the locality from which 
the type came must remain an open question. 


Specimen (0). 
The card is marked ‘‘Co-type, Chiton hirtosus, (Perony 
Blainville. M. M. Peron et Lesueur, 1803, I. King, M®> 886 


= Acanthopleura spinigera, Sow.’ On back it has the fol- | 


lowing notes, ‘‘Ile King, Chiton hirtosus, Peron, 233,’’ im 
Peron’s handwriting; two words that look like ‘‘Leplus 
grand, A. aculeata, L., I. King,’’ in Lamarck’s hand- 
writing, and ‘‘Acanthopleura spimgera, Sow., Thiele det.” 
This specimen is similar in sculpture and spicules to specimens 
in my collection from Port Darwin which I have considered 
are referable to Blainville’s Chiton gemmata. The shell is 
a good deal curled but is in good preservation and measures 
51x38 mm. This could not have come from Ile King, but 
possibly Baudin sailed north as far as Shark Bay, where this 
Acanthopleura occurs. Is it not possible that this is the 
missing type of Blainville’s Chiton gemmatus? Up to the 
present I have not been able to refer to the original descrip- 
tion of that species. 


581 


Specimen (c). - 

On another -card is a specimen, which I am calling (c), 
marked ‘‘Liolophura hirtosa, Peron; collected by Peron et 
Lesueur, 1803.”’ In Dr. Lamy’ S opinion this specimen is the 
black variety described by Blainville, 1825, as variety V. of 
his Chiton gemmatus. The shell shows very little sculpture, 
the dorsal area is eroded, but the rest of the shell is well pre- 
served. There are very deep growth-lines and ridges, which 
are only subpustulose in the lateral areas. It is curled and 
measures 30x23 mm. This spm.=Liolophura hirtosa, 
(Peron) Blainville. 

Note:—Blainville states that his variety V. was in the 
collection of the Paris Museum, but that the type of normal 
C’. gemmatus was in his own collection. 


Specimen card. 


This has two specimens mounted on it; they are marked 
“L. georgiana, Q. et G., Port du Roi George. ’ These are 
not that species, but are the Sydney shell LZ. gaimardi, Blain- 
ville. There are sufficient of the girdle spicules left to assure 
the correctness of the determination. 


Specimen (¢). 


This is in spirit and marked ‘‘Acanthopleura quatre- 
fagesi, Rochebrune (Rochebr., Bull. Soc. Philom., 1880-81, 
p. 117; Jour. de Conch., 1881, p. 44).”’ This is Liolophura 
hirtosa, (Peron) Blainville, and very probably was one of 
Blainville’s original shells. 

Liolophura (Chiton) see guana, Quoy et Gand Type 
(Voy. de ]’Astrol. Zool., 111., p. 379, t. 75, f. 25-30, 1833), 
Port du Roi George. There ‘are four specimens quite typical 
of this common Western Australian shell; as Peron’s name, 


_ hartosus, was published by Blainville in 1825, that name 


replaces that of Quoy et Gaimard. There are old labels 
attached reading, ‘“‘Chiton magellanicus, Chem.; Chiton 
georgianus, Q. et G. Type figured. Port du Roi George, 
New Holland, Expedition d’Urville, 1824, the figure in. Voy. 
Astrol., pl. 75, figs. 25-30, agrees with these specimens.’ 


Liolophura (Chiton.) gaimardz, Blainville (Dict. Sci. Nat., 


_ xxxvi., p. 546, 1825). The type was collected at Port J. ackson 


by Quoy and Gaimard and was preserved in spirit. This 
bottle contains two specimens with a more recent label, 
“Acanthopleura magellanica, Chem.” These may be the types, 
as the type is referred to as being in the Paris Museum in the 


_ catalogue of that Museum, dated 1838. 


582 


Onithochiton (Chiton) wndulatus, Quoy et Gaimard. 
Type (Zool. del’ Astrol., p. 393, t. 75, f. 19-24, New Zealand). 
The label is in the handwriting of Quoy or Gaimard, ‘‘P]. 75, 
figs. 19-24, 1833.’’ This corresponds with specimens in my 
own collection from Doubtless Bay, New Zealand, except that 
in the type the diagonal rib is almost smooth, showing little 
granulation: The shells are bleached. 


Omthochiton astrolaber, Rochebrune. Type (Bull. Soc. 
Philom., Paris, 1880-81, p. 120), New Zealand, Quoy et 
Gaimard, 1829. This shell has spaced granules in the diagonal 
rib similar to my Doubtless Bay specimens, and is only 
a slight variation from the type of Quoy and Gaimard’s 
undulatus. 

Onithochiton neglectus, Rochebrune. Type (Bull. 
Philom., Paris, 1880-81, p. 120), Wellington, New Zealand, 
Quoy et Gaimard. This is an exceptionally granulose shell, 
probably a variety. of Quoy et Gaimard’s wndulatus, but as 
that name was preoccupied Iredale substituted the name 
neglectus, Rochebr. (Trans. N. Z’d. Inst., vol. xlvii., 1914). 


Onithochitow lyelli, Sow. There is in spirit a rather 
worn specimen from Ile Pitcairn. This seems conspecific with 
O. quercinus, Gould. 

Gymnoplax adelaidensis, Quoy et Gaimard, 1829. This 
is an East Indian shell from Amboine. It has scales like a 
Haploplax and resembles members of that genus in general 
shape, but there the resemblance ends, the valves being very 
strongly sculptured. I have no reference to its description. 


583 


ECOLOGICAL NOTES ON SOUTH AUSTRALIAN PLANTS. 
PART 1. 


By Ernest H. Isine. 
[Read October 19, 1922.} 
Puates XXXVITI. ro XLII. 


I. INTRODUCTION. 


These notes are the result of a trip taken along the 
Transcontinental Line between Hughes and Kingoonya from 


_ September 5 to 24, 1920. Collections of plants were made at 


_ the following places showing the number of miles from Port 


— 


<< 


~ me 
a 
i - _ 


Augusta :—Hughes, 567 miles; Ooldea, 427 miles; Immarna, 
407 miles; Barton, 376 miles; Wynbring, 321 miles; Tar- 
coola, 257 miles; and Kingoonya, 209 miles. 

The rainfall over the area collected had been heavier that 
year than for a number of years, resulting in splendid growth 
of native vegetation. Seeds that were dormant for a 
number of years must have germinated that year, for there 


_ was.an abundance of plants at all the places visited. 


Reference will be made in this paper to the ecological 


_ factors noted in connection with the plants seen and collected 


at the various places mentioned. Plants were collected up to 
three miles from the centres referred to. 


Throughout the trip I was helped very considerably in 
collecting and drying by Mr. A. M. Lea, F.E.S., Government 
Entomologist, who was collecting insects for the Museum on 


_the same trip. 


An asterisk denotes an introduced plant. These were not 
seen to any extent and only close to the railway stations. 


For assistance in identifying some of the specimens I am 


“indebted to Mr. J. H. Maiden, I.8.0., F.R.S., F.L.S., etc., 
‘Director Botanic Gardens, Sydney (Hucalyptus and Acacia), 
Mr. J. M. Black, and Professor T. G. B. Osborn, D.Sc. 


II. PuysrtoGRAPuHy. 
1. THE NULLARBOR PLAIN. 
Size.—The Nullarbor Plain commences at Ooldea at its 


_ eastern boundary and stretches away westward to the border 


for 170 miles, and thence into Western Australia. Its 
_ Southern boundary is the coastline of the Bight, and it 


extends for about 100 miles north. 


m) a - | 


584 


The Little Plain.—At 441 miles from Port Augusta a 
ledge is met with which is the edge of the Nullarbor Plain 
proper. This is 17 miles west of Ooldea, and it forms a 
“little plain’’ which is quite distinct from the big plain 
further west. This small area is of an undulating character 
and grows a number of trees and small shrubs which appear 
to frequent the depressions. On the Nullarbor Plain, itself, 
this bigger growth disappears. It is on this small strip of 
country that the florulas of the plain and the sandhills meet, 
but there is very little invasion by the different plants on the 
neighbouring territory. 

The Plan Proper.—The Nullarbor Plain stretches away 
north, south, and west from the ‘“‘ledge’’ in an unbroken 
expanse of level, or slightly undulating, country as far as the 
eye can see. From the “‘ledge’’ (441 miles) to Hughes (567 
miles), which is within 32 miles of the Western Australian 
border, the country is the same uninteresting plain not relieved 
by any prominence whatever. Slight undulations occur, in 
places, and are from a quarter to half a mile, or more, across; 
but the resulting rises and depressions would only be about 
4 ft. or 5 ft. above or below the surrounding level. The 
rises, generally, have an outcrop of limestone with weather- 
worn fragments of the same lying around. The top soil, held 
together by the plants, is a reddish, friable, sandy loam which 
extends for at least 12 in. below the surface. In places it is 
of a clayey nature. In the depressions there is no surface 
limestone. These shallows (one large one at Hughes is called 
‘“The Dry Lake’’) grow fewer plants than the higher levels, 
and in them the ‘“‘Australites,’’ or ‘‘Obsidian bombs,’ are 
more readily found. The hollows do not hold water long. 
While we were at Hughes an inch of rain fell in one day, but | 
there was no water in the ‘‘lake’’ next day. : 

Several ‘‘blowholes’’ were seen at Hughes. They were 
about 15 ft. deep and about 3 ft. wide, with limestone ledges 
forming the sides. The bottom was soon reached by dropping 
a stone down, and no movement of air was observed going in 
or coming out. 


2. THE SANDHILLS. 


The sandhills commence at about 324 miles from Port 
Augusta, where they leave the stony undulating country. The 
sandhills are small at first but increase in size until some of 
them are 30 ft. to 40 ft. high and run in ridges for long 
distances. These ridges trend in almost every direction. At 
Ooldea they are east and west, and north-east and south-west. 
At Barton they are about east and west. The sand is fine 
and chiefly pinkish in colour; at Ooldea Soak, three miles 
north of the railway station, where the sandhills are very big, 


~ 


585 


the sand is almost white. At about 15 ft. below the surface 
at’ the Soak a very light-coloured clay is reached. This clay 
is very stiff and forms an impervious bottom for the wells 
that have been sunk. The wells are not sunk lower than the 
above depth and are timbered all the way down. The water 
soaks in within a few hours to about 3 ft. of the surface. 
There are eighteen wells at this spot and they are situated in 
a hollow surrounded by high sandhills. Twelve of them pro- 
duce beautiful, fresh, drinking water, while the other six are 
fit for human consumption but slightly brackish. 


There is no doubt this fresh water has been known to 
the natives for many miles around for generations, as native 
flint chippings can still be picked up in handfuls around the 
wells. It is a veritable oasis, and has been made use of by 
early explorers. The sandhills are clothed with a dense vege- 
tation comprising trees (up to 40 ft. or more in height), 
shrubs, undershrubs, small perennials and annuals. A fine view 
was obtained from the top of a tall sand ridge at the Soak, 
and the prevailing mallee sandhill scrub stretched away to the 
north, east, and south as a dark expanse of country. 


For most of the year the plants of the sandhills are 
subjected to very severe growing conditions, and transpiration 
must be at its maximum during that period. Such conditions 
tend to keep an open formation; that is, plants have open 
spaces between them of some yards. Yet often in the hollows 
between the sand ridges the Acacias and other shrubs are so 
close together that they touch one another, and one has to 
push a way through them. The vegetation has responded to 
its environment by developing narrow leaves (or phyllodes 
in the case of the Acacias), thus reducing transpiration to a 
minimum. The broad-leaf plants, such as Eucalypts (L. 
oleosa, EH. pyriformis, and EF. transcontinentalis), have re- 
sponded to the prevailing meteorological and edaphic factors 
by producing coriaceous leaves with few stomata which are 
deeply set below the epidermis. The small herbaceous annuals 
grow chiefly out in the open, it was rare to find them growing 
below the larger shrubs or trees. The annuals consisted largely 
of composites, although Calandrima polyandra, the “‘para- 
keelya,’’ formed large patches around Barton. 


The sandhills are fixed, being clothed with native vege- 
tation. When the covering is removed trouble is experienced 
with drifting sand. This has been the case in some of the 
railway cuttings, which have had to be faced with a retaining 
mat consisting of stakes, boughs, and small branches. 


The sandhills grow a greater number of plants than any 
other portion of the country visited along the line. - : 


586 


3. THE COUNTRY AROUND TARCOOLA AND KINGOONYA. 

At about 324 miles from Port Augusta, near Wynbring, 
the sandhills disappear and an undulating stretch of country 
is entered upon, which continues to Kingoonya. At Tarcoola 
there are some small hills, the sloping sides of which are 
thickly strewn with rock fragments, about 4 in. square. 


Tif. Prants oF THE NULLARBOR PLAIN. 

General.—There are two main types of plants at Hughes: 
(a) shrubs of about 50 cm. in height, and (>) small herbs 
and grasses. This formation was constant, as far as observed, 
for 140 miles between Ooldea and Hughes. It was the result, 
no doubt, of the uniform character of the surface topography, 
soil, and rainfall. The shrubs include a very few tall ones of 
Pittosporum phillyracoides and Acacia tetragonophylla, and 
it is a remarkable fact that there are so few of them. 

The plants may be considered according to their height. - 

1. The tallest plants were shrubs, 2 to 3 m. in height, 
consisting of “‘dead finish,” Acacia tetragonophylla (only one 
plant seen, 3 m. in height), and the ‘‘Weeping Pittosporum,’’ 
P. phillyraeoides, of which only a few shrubs came under 
notice. 

2. The bluebush and saltbush shrubs varied from a half 
to one metre in height, and were the dominant shrubs of this 
vast treeless, riverless plain. 

3. The undershrubs and larger perennials and annuals, 
of from 20 to 45 cm. in height, formed this third range of 
plants, and consisted of species of Aochia, Bassia, Blennodia, 
Swaimsona, composites and grasses. 

4. The ground flora of only a few inches in height was 
represented by composites, and by Calandrima, Daucus, 
Hrodium, Euphorbia, Lepidiwm, Lotus, Nicotiana, Plantago, 
T'etragonia, Crassula, and Zygophyllum species. This arrange- 
ment, however, does not give the ecological relationships which 
I wish to emphasize. 

The following formations, which are of the open type, 
were noted on the plain. 


Bluebush Formation. 
IPLSxl SS fie 1: 


The Nullarbor Plain is not a dead level, but consists of 
undulations, forming slight rises and shallow depressions, 
varying from 1 to 2 metres. The bluebush (Kochia sedifolia) 
was not confined to either the rises or the hollows, but it was 
noted that this shrub dominated an area of several hundred 
square yards in extent. The saltbush (Atriplex vesicarvwm ) 


587 


was not completely excluded from this region, but the blue- 
bush gave it a characteristic blue-grey appearance. A very 
prominent species in this formation is Goodenia pinnatifida, 
which covered numerous areas and ranged from 10 to 12 cm. 
high. It was in full flower at the time of my visit (Sep- 
tember 8) and was a beautiful sight. Podolepis canescens was 
found in this station, and it is a larger plant than the pre- 
vious one but not nearly so plentiful: It brightened the dull 
hue of the bluebush foliage. 

To be seen in some numbers with the above plants was 
an interesting variety of Calotis multicaulis (n. var. brevi- 
radiata, see p. 604), a small diffuse herb 5 to 20 cm. in’ 
height. It was growing in little colonies of about a metre 
across. Another plant growing chiefly in colonies, but much 
more plentiful than the last species, was Cephalipterum 
Drummondu, a species with dense white heads. Some speci- 
mens collected were remarkably small, being only 34 cm. in 
height, while the largest were 15 cm. _ 

The following plants were often found in association, 


usually in small depressions in which water remained for a 


short period after the rain :— 

Helipterum strictum, growing up to 25 cm. in height 
and dominating the association. Vuttadima australis, in 
lesser numbers and not so high. Daucus glochidiatus, about 


_ 20 cm. in height. Podocoma nana, plentiful, but only up 


to 8 cm. in height; this is the first record of this plant for 
Nullarbor Plain. Cvassula Sieberiana, varying from 3 to 
6 cm. Tetragonia expansa, a plant quite prostrate and 


_ spreading 20 cm. or more. Plantago varia, the smallest plant 


in the colony, being only 3 cm.(?) or less in height. And 
Calandrima pusilla, another small annual. 

A small sticky composite (Heli:pterum tenellum) formed 
areas of several square feet ; the plants ranged from 6 to 18 cm. 
in height. Smaller still, and growing together, were Bassia 


_selerolaenoides and B. patenticuspis, which formed an open 


association. Two species of Zygophyllum (Z. iodocarpum and 


_ 4. ovatum) were associated and grew in considerable numbers 


where the ground was subject to flooding. A small composite 
(Minuria leptophylla) was not often seen, but (nephosis 


_skirrophora was much more plentiful. A dwarf annual cruci- 


fer (Thlaspi Drummondii) was fairly common in this station, 
as was also Lepidiwm rotundum, DC., var. phlebopetalum, 


_ Maid. et Betclie, a plant only 4 to 8 em. in height. The tiny 


annuals—Plantago varia, Calotis hispidula, and Tsoetopsis 


. granunifolia—were fairly numerous between the bluebush 
_ shrubs. A common composite was Vittadinia australis, and 


one, much less so, was Hlachanthus pusillus, and an annual 


588 


that grew in numerous patches was Siloverus brachypappus, 
which is a small diffuse annual of from 2 to 6 cm. in height. 
This latter plant was a notable feature in many places on the 
Nullarbor Plain visited. Two dwarf plants not often met 
with were Hrodium eygnorum and Convolvulus erubescens. 
At the time of my visit the most abundant plant, and the one 
which covered a large area, was the white everlasting Helip- 
terum floribundum. It is a very showy annual growing up to 
25 cm. 

The wooliy bluebush (Aochia villosa) was a rare plant 
on the Nullarbor Plain, as also was A. Georgei; they were 
smaller plants than the typical bluebush (Kochia sedifolia). 
Growing among species of Zygophyllum were plants of 
Lepidium papllosum. The introduced pest *Hmex australis 
was spreading in the open spaces near the railway line at 
Hughes. Another rare plant was Lepidium fasciculatum, but 
was more plentiful around Tarcoola. Two plants found in 
open association were Swainsona Oliveri and Sida corrugata, 
var. orbicularis. Among the rare species were noted Salsola 
Kah, var. strobiifera, Senecio brachyglossus, Huphorbia 
Drummondii, and Minuria Cunminghami. Three grasses were 
identified: Stipa eremophila and S. scabra, var. auriculata, 
and the dwarf Danthoma penicillata; the two former were 
much more plentiful than the latter. 


Saltbush Formation. 


The saltbush (Atriplex vesicarium), like the bluebush, 
is a perennial shrub of about 60 cm. in height. Usually it is 
just a little shorter than the bluebush, and the two species 
form the main vegetation of the Nullarbor Plain. 

On the whole, the species observed in association with the 
bluebush were also noted among the saltbush. There were, 
however, certain plants only seen with the saltbush. In 
depressions there was less vegetation than on the higher 
ground ; the smaller plants (annuals chiefly) were absent, and 
the formation was decidedly an open one. It was in this 
station only that the following plants were seen :—Hremo- 
phila maculata, a shrub about 45 cm. in height; Atriplex 
campanulatum, a small saltbush 25 cm. in height; spear 
grass, Stipa eremophila (also observed in the bluebush forma- 
tion); Blennodia trisecta; and the decumbent plant, 
Frankema paueiflora. The annuals were of few species and 
sparsely distributed, including Lotus australis, var. parvi- 
florus, a plant with prostrate stems and often spreading to 
1 m. across; Lavatera plebera, of about 30 cm. in height; 
small plants of Micoteana suaveolens; and Swainsona 
phacoides, often wide spreading. | 


589 


The Plain and Sandhills. 


There is not much change in the general aspect of the 
flora where the Nullarbor Plain joins the sandhill region. 


Just before leaving the ‘‘Plain’’ the “dead finish” (Acacia 


tetragonophylla) becomes more plentiful, but it was seldom 
seen in the sandhill country. The following plants were noted 
just west of the sandhills and were common to both types of 
country :—(Goodenia mnnatifida, Cephalipterum Drum- 
mondir, Calotis hispidula, Kochia sedifolia, Stipa scabra, var. 
auriculata. There is very little overlapping of the plants of 
the two regions. 
IV. Tue Sanpaiuts Fora. 


1. OOLDEA DISTRICT. 


The sandhills’ flora is of a typical sclerophyllous nature, 
and here again the formation is of an open character. 

T'rees.—The trees and larger shrubs usually have reduced 
leaf surfaces. In the case of Casuarina lepidophloia the leaves 
are represented by very small sheathing teeth, and the branch- 
lets are only 1 mm. in width. Myoporum platycarpum was 
sparingly distributed, and much less so was Heterodendron 
oleaefolium, both of which have flat leaves. The latter was 
usually found with Acacia ramulosa in the flats between. the 
sand ridges. The mallees were not so plentiful as the wattles, 
and two of the broad-leaved Eucalypts were FH. oleosa and 
EL. transcontinentalis (pl. xxxix., fig. 2), which formed the 
bulk of the mallees. #. pyriformis seemed confined to a small 
patch at Ooldea Soak. Amoag other mallees were /. 
uncinata and EF. gracilis, forming large shrubs and growing 
interspersed with Acacia ramulosa. The quandong (Fusanus 
acuminatus) was not common and seemed to prefer the sand 
ridges. 

Shrubs.—The phyllodes of some of the acacias were nar- 
row and hard, such as A. tetragonophylla, A. colletiowdes, A. 
ramulosa, and A. aneura, the last two being more plentiful; 


while A. Randelliana and A. Burkittii were not seen to any 


extent. Of those with broader phyllodes A. Kempeana, A. 
Osswaldii, and A. ligulata were fairly numerous. Other 
shrubs were Hremophila alternifolia, which was seen in fair 
numbers and often associated with Casuarina lepidophloia; 
Eremophila Latrobe: and its variety Tietkensi were the next 
most plentiful, but 2. Gibsonw was rare. Two grevilleas (G. 
pterosperma and G. stenobotrya) were usually seen growing 
on the flat ground, but of infrequent occurrence. The para- 
sites, Loranthus linophyllus and L. pendulus, were somewhat 


rare, the former growing on Heterodendron oleaefolium and 


the latter on Fucalyptus transcontinentalis. Rhagodia 


590 


Billardieri formed tall shrubs and was often seen among 
acacias. In the wide open flats, between some of the smaller 
sand ridges, were seen shrubs of Cratistylis conocephala and 
Westringia Dampieri, var. rigida, and Bassia echinopsila 
was associated with them. Two other shrubs, Cassia eremo- 
phila and C. Sturtw, were fairly common in the sandhill 
region, with Dodonaea attenuata as a rare species. 

The Ground Flora.—By the ground flora is meant the 
undershrubs and annuals which range from 2 to 25 cm. in 
height. The dominant species were Cephalipterum Drum- 
mondu, Waitzia acuminata, and Helipterum floribundum; 
they are annuals and grow in open association. Some plants 
preferred the sand ridges (often in the open and seldom in 
the shelter of other larger plants), wiz., Waitzia acuminata, 
Calandrinia disperma, Stackhousia muricata, Podotheca 
angustifolia, and Pomaz umbellata. The latter species. and 
the Stackhousia, developed a long slender tap-root which, 
no doubt, penetrated the loose sand to the moisture below; 
the lateral rootlets were not robust, as the plants depended 
on depth of root rather than on spread. The poor rainfall 
(see table) of the district and the intense heat, combined 


RAINFALL FOR 


} 
1914 | 1915 | 1916 1917 1918 | 1919 | Aug.| Average 


31,1920 
Hughes peal pee il aa aut lc — _\. 6 Toes — 
Ooldea i an — |7:35) 6°75 | 4:29} 7-05*(2) 
Immarna sale —— ee | — — ~ | 6°65 | 5°14 — 
Barton ee sey — | — 1/763} 6-44 | 4:35] 7-03*(2) 
Tackpale _..,5-28t) 3°76 | 7-92] 9-20] 7-49 | 7.35 | 5-87 | 7 33417) 
Kingoonya . |) a= | 8-00 | 8°65 | 5°76 | oP Gy 7-05*(4) 
Eucla ... .«| 9°00 12°77 |{9'51 | 10°70 5°18 |10-05+(44) 


/ 


Port Augusta .. 8°80 | 11.26 | 13°67 | 7°58 | 967 7°76 | 9°43*(60) 


bedi Lt 
| 


* () Number of years for average 
+ These totals are shown in inches 


with an extreme evaporation, tends to the production of an 
elaborate root system. This is specially necessary in the 
plants growing along the tops of the sandhills. Growing 
chiefly on the flats, between the sandhills, were: Zygophyllum 
fruticulosum, Euphorbia Drummondii, varieties of Sida 


— — ———— 
. ‘ . 


I gg i OI AE 


Oe = 


Ft? a0" 


591 


corrugata, in open association; while Uldinia mercurialis, 
Lappula concava, Calotis hispidula, and Daucus glochidiatus 
formed patches often in association with one another. Also 
on the flats, Velleia paradoxa was found associated with the 
annual plant Brachycome ciliaris. 


2. OOLDEA SOAK. 
At Ooldea Soak, where a wonderful supply of fresh water 
is obtainable at a shallow depth, some of the vegetation is 
luxuriant; for instance, Myriocephalus Stuartii formed a 


_ veritable carpet where it grew in the hollows near the wells. 


— 


Associated with this plant was Senecio Gregorw and large 
shrubs of Leptospermum laevigatum, var. minus (pl. xxxix., 
fig. 2), although I also noted the latter species some miles 
from the Soak growing on a sand ridge. In the hollow, 
where the wells are situated, was found the ‘‘water-bush, 
Adriana Hookerit, and ascending the sandhills, to the west, 
Melaleuca parviflora and Acacia ligulata were met with, 
while Gyrostemon ramulosus was only seen on the ridges. 


3. BARTON DISTRICT. 


Barton is situated in the centre of the sandhill tract and 
is similar country to Ooldea. Its flora, too, is similar, only 
slight differences being noted. Twenty-six of the species noted 
here were not recorded from Ooldea, while 67 species col- 
lected at the latter place were not seen at Barton. The type 
of plants was the same as at Ooldea, Casuarina lemdophloia, 
however, was more plentiful, although it could hardly be said 
to dominate the flora. There was the usual A cacia-Eucalyptus 
association with Acacia ramulosa and Eucalyptus oleosa, as 
the dominants, particularly the former. The former species 
was met with almost everywhere (sometimes in. a semi-closed 
formation), while the latter was reduced to a clump, here and 
there. Although the season (1920) had been a good one 
hardly a seedling was seen of either of these species. An 
occasional clump of FH. transcontinentalis was seen, while a 
clump of mallee (Hucalyptus oleosa, pl. xli., fig. 1), remark- 
able for its prostrate trunks, covered a patch about 10 yards 
across, situated in a hollow between the usual sand ridges. 
Only two or three of the trunks were upright and were about 


3m. high; the others were lying on the ground, right from 


their base. The middle of the trunk was somewhat arched 
and the branches were horizontal. The aphyllous shrub, 
Bossiaea Walkeri, seemed to prefer the lower situations and 
often formed large thickets. The flattened stems exude quite 
a quantity of smooth white powder while drying. Dodonaea 
microzyga was not plentiful, nor was Olearia subspicata, and 


592 


both grew on the flats with Casuarina, Grevillea Huegelii, 
and Acacia colletioides. Hremophila scoparia was associated 


with Cassia eremophila and C. Sturtii, and, in places, formed | 


quite a distinct feature of the vegetation. Vhryptomene 
Eliott was seen on a sand ridge at Barton. 

In a photograph (pl. xli., fig. 2) taken at Barton 
Thryptomene LEiliottw is. seen in the foreground with 
Casuarina lepidophloia and Hucalyptus close by. In another 
situation, Casuarina leyidophioi is growing with acacias, 
mallee, and Triodia irritaws. This latter plant was fairly 
common at Barton, and, in another place, it was noticed 
associated with Solanum coactiliferum, Acacia ligulata, and 
Thryptomene Elhottu. The common ‘‘parakeelya’’ (Calan- 
drima polyandra) of the sandhills was growing so profusely 
in places that it became almost a closed formation, its asso- 
ciates in one place were Helichrysum lucidum and Solawum 
orbiculatum. In open, flat ground Trichinium corymbosum 
and Podolems camllaris were associated; they are both small 
annuals. 


V. DESERT FORMATIONS OF THE TARCOOLA REGION. 
CLIMATIC AND EDAPHIC FACTORS. 


The sandhill region is left at Wynbring, where, travel- 
ling east, an undulating stony country is entered upon. As 
was to be expected, the flora changed as soon as the sand- 
hills were left behind. The vegetation now was not so dense 
or plentiful, no doubt caused by the dry subsoil. The top 
soil is of a clayey nature in this region and surface water 
would remain longer than in the sandhills. In the sandy 
country more moisture reaches the subsoil, which proves to 
be of a wonderfully retentive nature; there is, consequently, 
a greater amount of moisture available for the plant cover- 
ing. This influences the flora of the two regions under 
discussion. 

KOCHIAS AND ACACIAS OF TARCOOLA. 


The predominating species in this station is Aochia 
sedifolia and Acacia Loderi, while Kocha triptera and 
Eremophila rotundifolia are represented by numerous plants. 
Also Acacia aneuvra claims attention, as it was frequently seen ; 


Hakea leucoptera was not so plentiful. The plants on top of 


a rocky hill (pl. xlii., fig. 1) consisted of Acacia tarculensis 
and Trichinium iwcanum, which were the dominants; here 
and there Rhagodia Gaudichaudiana and Hnchylaena 
tomentosa were seen, while the smaller plants, Helpterum 
Fitzgibbon and H. pterochaetum were fairly numerous. : The 
rocky slopes of the low hills have a distinctive flora, and, 
besides the prevailing bluebush, Helipterum Humboldtianum, 


% 


Se —= » a . ry —— 


593 
is seen along a small dry watercourse, and associated with it 
is H. moschatum. Larger plants here were Sida calyxzhymenia 
and Atriplex vesicarium. 

Where the slopes led into more flat country Kochia 
pyramidata and Clianthus Dampier: were associated, and, in 
open formation with the bluebush, the following plants were 
noted :—Kochia villosa, Bassia diacantha, B. paradoxa, and 
Salsoli. Kali is only represented by its variety strobilifera, and 
it was rare here, as it was on the Nullarbor Plain. 

Coming right down to the depression at the base of the 
hill near Tarcoola, the succulents, A2z0o0ow quadrifidum, form- 
ing small shrubs, and Tetragoma expansa, were in association 
with a few plants of Zygophyllum Billardiert, var. ammo- 
philum, with them. 

On the extensive clay flats the tall Acacia aneura was 
the dominating species, and Calogyne Berardiana was also 
very plentiful, and formed large patches in places with asso- 
ciations of Goodenia pinnatifidia and G. pusilliflora. Grow- 
ing in the shelter of the former were A butilon oxycarpum 
and Huphorbia eremophila. In this formation was also seen 
Brachycome ciliaris, Cephalipterum Drummondii, Lepidium 
rotundum, var. phlebopetalum, Calatis hispidula, Stewo- 
petalum lineare, Erodium cygnorum, and Millotia tenurfolia, 
the first species being the most plentiful. 

Where the soil was of a more sandy loam in this area 


_ the vegetation was more pronounced. Several patches of this 


nature were seen, and the predominating plants were 
Helichrysum Lawrencella, var. Davenportu, Craspedia 
plerocephala, Myriocephalus Stuartiz, Swainsona phacordes, 
and S. microphylla, the last two specially so. A few plants 
of Calotis multicaulis, Templetonia egena, and Helipterum 
floribundum were not so common. In a small depression, 
Frankema serpyllifolia had almost made a closed formation. 
No eucalypts were seen at Tarcoola, but scattered species of 
Eremophila were noted as follows:—F. Duttonu, EF. glabra, 
EF. latifolia, E. Latrobe, and EF. Paisleyi, besides those 
already mentioned. : 


KINGOONYA PLAIN. 


The dominant species of the plain was Acacia Loderi, 
and with it was associated Minwria leptophylla (pl. xli., 
fig. 2). The shrubs noted were Rhagodia Gaudichaudiana, 
Koch triptera, K. villosa, Cassia Sturtu, EKremophila alter- 
nifolia, HL. Latrobei, and Bassia paradoxa. Smaller plants 
were Bassia sclerolaenoides, Rutidosis helichrysoides, Ixiolaena 
leptolepis, and Leptorrhynchus tetrachaetus, var. penicillatus, 
and were only represented by few specimens. In other forma- 
tions were Blennodia trisecta, Menkea australis, Chanthus 


594 


Dampieri, Podocoma nana, Gnephosis cyathopappa, Helip- 
terum Charsleyae, H. stipitatwm, and other small annuals. 

Where the ground was lowlying the grass Hragrostis 
Dielsii was recorded, and with it were Swainsona Oliveri, 
Tribulus terrestris, Zygophyllum fruticulosum, Z. ovatum, 
and Isoetopsis gramimfolha. 


VI. A CENSUS OF AND Notes on PLANTS COLLECTED. 

References: —H., Hughes; O., Ooldea and Ooldea Soak; 
I., Immarna; B., Barton; W., Wynbring; T., Tarcoola; K.; 
Kingoonya. The numbers following the capital letters refer to 
my specimen number. The above places are in Tate’s District W, 
as shown in Tate’s ‘‘Flora of Extrat. South Australia,’”’ p. 204. 
Where a plant is new for this district, ‘‘Dis. W.’’ is shown. An 
asterisk denotes an alien species. 


POLYPODIACEAE. 
Cheilanthes tenuifolia, Swartz. T. 1726. 


MARSILIACEAE. 

Marsha Drummondu, A. Br. K. 1846. Appears to be 
this species, although the sporocarps are very shortly stalked 
(3-4 mm.) and the cases are about the same length, hairy, 
and with a few oblique transverse ridges. Leaflets ovate- 
cuneate, hairy, but becoming glabrous with age. Near J. 
hirsuta, R. Br. 


SCHEUCHZERIACEAE. 

Triglochin centrocarpa, Hook. O. 1609, T. 1799. 

My plant No. 1609 agrees well with the illustration (pl. 
iv., 2) by Ostenfeld in Dansk Bot. Arkiv., Bd. 2, Nr. 8, 1918, 
but they are taller, 7.¢., 13 cm. high. The flowers are dis- 
tanctly pedunculate. 

The Tarcoola specimen (No. 1799) is only 3°5 cm. high; 
- the fruits are as long as No. 1609, but the spur is more 
pronounced. 

GRAMINEAE. 


Identified by Mr. J. M. Black. 


Panicum leucophaeuwm, H. B. et K. T. 1648. 

Pappophorum nigricans, R. Br. T. 1642, 1644. 

Stipa eremophila, Reader. H. 1636. 

S. scabra, Lindl. W. 1216. 

S. scabra, Lindl., var. auriculata, J. M. Black. H. 
1637, O. 1640. 

Aristida stipoides, R. Br. T. 1643. 

Danthonia pemcillata, (Labill.), F. v. M. H. 1638-9, 
K. 1649-50. 

Diplachne loliiformis, F. v. M. K. 1647. 


. 595 


Triodia writans, R. Br. O. 1293, I. 1246, B. 1317. 
Eragrostis Diels, Pilg. K. 1646. 

E. eriopoda, Benth. T. 1411. 

E. eriopoda, Benth.; var. lamflora, J. M. Black. O. 1641. 


LILIACEAE. 
Thysanotus extliflorus, F. v. M. O. 1625. Petals in my 
specimens are not minutely fringed, but it agrees 
with the above otherwise. 


CASUARINACEAE. 


Casuarina lepidophloia, F. v. M. O. 1479, B. 1705. 
Teeth, 8 or 9; cones, 12 to 15 mm. long. 


URTICACEAE. 


Humulus, sp. O. 1282. A single specimen growing near 
the ballast on the railway line. 


PROTEACEAE. 
Grevillea Huegelu, Meisn. B. 1339. 
G. pterosperma, F. v. M. O. 1302, B. 1383. 
| G. stenobotrya, F. v. M. O. 1302a. In bud only, Sep- 
| tember 16, 1920. 
Bukea-leucopiera, BR, Br... 'T; 1785... ““Dis: .W.7 


SANTALACEAE. 
Fusanus acuminatus, R. Br. O. 1614, K. 1832, B. 1706. 


Loranthus linophyllus, Fenzl. O. 1589. Growing on 
Heterodendron oleaefolium, Desf. 

L. pendulus, Sieb. O. 1289. Of pendulous habit grow- 
ing on Hucalyptus transcontinentalis, Maiden. 


} 
: LORANTHACEAE. 


POLYGONACEAE. 
*Emex australis, Stem. H. 1547a, T. 1777. 


CHENOPODIACEAE. 
Atriplex campanulatum, Benth. H. 1508, O. 1602. 
BD Sg oad 
A. spongiosum, F.v. M. B. 1360, K. 1807. 
A. vesicarvum, Hew. H. 1228, 1259, 1260, 1511, 1541, 
; 154va, 1o65ay W. 1395, 7.1714, 1761. «The fraits 
| of this species vary a good deal. In one specimen 
from the Nullarbor Plain (No. 1565a) the append- 
ages have thick prickle-like lobes covering them. In 
No. 1761 the fruiting calyx is entire, semi-orbicular, 
and with very small appendages; the leaves are 
small, mostly orbicular-ovate. 


. 


596 


Rhagodia Billardiert, R. Br. O. 1604. 
R. Gaudichaudiana, Mog. O. 1620, B. 1351, W. 1202, 
T. 1408, K. 1836. 

Chenopodium cristatum, F. v. M. O. 1701, B. 1384. 

Enchylaena tomentosa, R. Br. B. 1331, T, 1778. 

Kochia George:, Diels. H. 1226, 1542-3, T. 1723, 1756. 
First record for Nullarbor Plain, Tarcoola, and for 
the State. Originally described from Western Aus- 
tralian specimens by Diels and Pritzel in Bot. Jahrb., 
184, 1904, with a figure of fruit (fig. 20, D). Pre- 
viously confused with glabrous forms of A. villosa, 
but the obconic base of the fruiting perianth is a 
very distinct feature. 

. pyramidata, Benth. T. 1788. 

. sedifolia, F.v. M. H. 1227, 1258, 1544. T. Neither 
in flower nor fruit. 

. triptera, Benth. T. 1763, K. 1829. 

. triptera, Benth., var. erioclada, Benth. O. 1235, 
B. 1380, T. 1724, 1762. 

. villosa, Lindl. O. 1286, T. 1722, 1742, K. 1833. 

assia biflora, F.v. M. K. 1805. ‘Dis. W.” 

. dracantha, Woy. MT. W018,. 8137; Tiare 

echinopsila, F. v. M. O. 1275, B. 1708. 

ertacantha, F. v. M. (B. lamcuspis, F. v. M.). 

Ao, 

sp.(?) O. 1284, 1603. 

. paradoxa, F. v. M. T. 1743, K. 1808. 

. patenticuspis, R. H. Anderson. H. 1230, 1513, 1548, 
O. 1285. This identification was made by Mr. J. 
M. Black, who advises that Mr. R. H. Anderson, of 
Sydney Botanic Gardens, is engaged on a revision 
of the Australian genus Bassia and has recently 
created this new species. 

B. sclerolaenodes,.F. vy. M.,..0;,1567, 1578, Te ties 

K. 1809>°)5 Diese RVs 
Pachycormia tenuis, (Benth.) J. M. Black. T. 1764. 


A RR RR 


mht a 


*SDigsy Wes 
Sdlsola Kah, ., var. strobélfera, Benth. H. 1569, 
T. 1793. 
AMARANTACEAE. 
Trichinium alopecuroideum, L. O. 1238, 1627, B. 1702, 
T.f72ik 


T. corymbosum, Gaud. B. 1311, T. 1798. 
T. exaltatum, Benth. O. 1287, 1626. 
T. incanum, R. Br. O. 1264, 1628, T. 1407, 1746. 


597 


PHYTOLACCACEAE. 
Gyrostemon ramulosus, Desf. O. 1303. 


AIZOACEAE. 
Tetragona expansa, Murray. H. 1536, T. 1732, 1186. 
Aizoon quadrifidum, F.v. M. T. 1760. 


PORTULACACEAE. 

Calandrima disperma, J. M. Black. O. 1588. 

C. polyandra, Benth. O. 1236, 1276, 1605, 1606, B. 
1232, 1385. Nos. 1276, 1605, and 1385 are the 
white-flowered variety. 

C. pusilla, Lindl. H. 1546, I. 1249, B. 1365, 1387, 
T. 1769. No. 1387. Plant larger than usual and 
more branching, stems 18 cm. long, racemes many- 
flowered. 

: CARYOPHYLLACEAE. 

Spergularia rubra, Camb. K. 1840. 


| PAPAVERACEAE. 
*Papaver hybridum, L. O. 1697. 


CRUCIFERAE. 

| Blennodia canescens, R. Br. W. 1393, T. 1717. 

B. curvipes, F. v. M. T. 1767. 

B. trisecta, Benth. H. 1257, 1549, K. 1810. 

*Sisymbrium orientale, L. O. 1698, T. 1195, 1196. Of 
the introduced species this one was the most common. 


Stenopetalum lineare, R. Br. O. 1262, B. 1364, T. 1748, 
K21839. °.“Dis:. W..’’ 

Menkea australis, Lehm. T. 1830, K. 1831. ‘‘Dis. W.”’ 

Thlasm Drummondii, Benth. H. 1520, 1700. A rare 
plant only collected on the Nullarbor Plain (Capsella 
Drummondu, F. v. M.). 

Legmdium fasciculatum, Thell. H. 1554, K. 1834. ‘Dis. 


mand ——E—E—— SS 


Ws’ 
L. pamllosum, F. v. M- H. 1593, O. 1547, T. 1201. 
i L. rotundum, DC. T.'1189. ‘‘Dis. W.’’ 


L. rotundum, DC., var. phlebopetalum, Maid. et Betche. 
B24 Os Taal yer: L290." “Dis. Woe : 


CRASSULACEAE. 
Crassula colorata, (Ness.) Ostenf. T. 1776a. 
C. Sieberrana, (Schult.) Ostenf. H. 1545, T. 1775. 


098 


PITTOSPORACEAE. 
Pittosporum phillyraeoides, DC. H. 1550. 


LEGUMINOSAE. 


Daviesia ulicna, Smith. B. 1367. It was quite a sur- 
prise meeting this plant, which is usually found in 
the Mount Lofty Range and the south-east of the 
State. Only one plant was seen, and was nearly 
2 metres high ; the bark was dark, rough, and ribbed. 
The flowers are in short axillary umbels, with the 
pedicels longer than the peduncle. ‘‘Dis. W.”’ 

Bossiaea Walkert, F. v. M. . B. 1217, I:.1244.-) fae 
flower September 5, 1920. Young branches silky | 
with dense adpressed hairs. 

Templetonia egena, Benth. T. 1729. 

Chanthus Dammert, Cunn. T. 1772, K. 1814. 

Swainsona Burkes, F. v. M. I. 1247. ‘‘Dis. W.” a 

S. microphylla, A. Gray. T. 1725, 1739: The leaflets 
vary a good deal in size and shape. I have them 
from 6 to 12 mm. long and from ovate to oblong. 

S. Olivert, F. v. M. H. 1563, K. 1841. 

S. phacoides, Benth. H. 1256, 1539, W. 1394, T. 1740. 
‘Diss Wet 

Psoralea patens, Lindl. K. 1835. 

Lotus australis, And., var. parviflorus, Benth. H. 1564. 
Flowers pink. 

Cassia artemisioides, Gaud. T. 1749. 

C. eremophila, Cunn. O. 1283, B. 1707. 

C. eremophila, Cunn., var. platy poda, Benth. O. 1272, 
1D 75, 1 608; %e 1416. 

C'. Sturin, R:; Br. _O. 1271, 1280, B. 1332, T. ATT 
K, 138i2. 


Acacia species identified by Mr. J. H. Maiden, I.8.0., 
F-R.S., ete 


Acacia aneura, KF. v. M. O. 1273, 1487, By iS2ee 
T. 1413, 1498. No. 1498 is a small intricate shrub 
of nearly 1 metre high. The branches are somewhat — 
angular with white scaly angles or lines; the phyllodes . 
are short and broad. Altogether the plant is very © 
different from the typical tree; this may be accounted — 
for by the fact that the only shrub seen was growing 
among rocks on the top of a rise near Tarcoola. 
Not in flower or fruit. 

A. brachystachya, Benth. T. 1414. 

A. Burkittu, F. v. M. QO. 1486. 

A. colletioides, Cunn. O. 1298, B. 1341. 


599 


A. Kempeana, F.v. M. QO. 1491, B. 1338(?). 

A. Loderi, J. H. Maiden. T. 1496, 1499, 1500, K. 1501. 
New for South Australia. Mr. Maiden described it 
in the Proc. Roy. Soc. N.S. Wales, vol. liii. (1920), 
p- 209, from Broken Hill specimens. It is a small 
tree 3-5 cm. high with branches and phyllodes fairly 
erect. The phyllodes vary from 25 to 90 mm. long 
and 1 to 2 mm. wide. Veins about 10, the central 
one on the surface of the phyllode is somewhat ridged. 
Pods almost sessile, light brown, 25-40 mm. long 
and 2°25 mm. wide. (‘‘Nos. 1500 and 1501 are more 
glabrous forms with narrower phyllodia.’’— 
hala, NE: } 

A. Oswaldu, F. v. M. O. 1494, B. 1345. 

A. Prann, J. H. Maiden.. B. 1330. New for South 
Australia. Mr. Maiden’s description is to be found 
in the Proc. Roy. Soc. N.S. Wales, vol. lh. (1917), 
p- 238, and was first collected near Kalgoorlie, Wes- 
tern Australia. It is a small shrubby tree of nearly 
3 m. high with spreading branches which start at 
the base of the trunk. The phyllodes are 25 to 
75 cm. long and 15 mm. wide, rigid, and spinescent. 

' Flowers in short axillary racemes. Pods not seen. 

A. ramulosa, W. V. Fitzg. O. 1268, 1269, 1274, 1489, 
B. 1333-5, 1492, 1495, 1502. New for South Aus- 
tralia. First described by W. V. Fitzgerald in Jour. 
W. Austr. Nat. Hist. Soc., No. 1, May, 1904, p. le, 
from specimens collected at Lennonville, Western Aus- 
tralia. A shrub with branches spreading from the 
base. Phyllodes up to 17°5 cm. long and 1-14 mm. 
wide, compressed terete. Veins many, very faint. 
Flowers yellow in cylindrical spikes of 12 mm. long, 
peduncle 12 mm. long. Pods 12°5 cm. long and 
4°55 mm. wide, somewhat constricted between the 
seeds, with longitudinal narrow strips of white and 
green. Very like 4. linophylla, W. V. F. 

A. Randelliana, W. V. Fitz. O. 1294. 

A. salicona, Lindl. O. 1488, 1493, I. 1248, B. 1313, 
1315. I do not agree with Mr. Maiden’s identifica- 
tion, but follow Mr. Black (Trans. Roy Soc. S. 
Austr., vol. xliv. (1920), p. 375,, and pl. xxiu., 
figs. 6 ‘to 11) in regarding this plant as A. ligulata, 
A. Cunn. 

A. tarculensis, J. M. Black. T. 1497. 

A:. tetragonwophylla, F. v. M. H, 1490, O. 1485, iBa13390. 

*Medicago denticulata, Willd. O. 1629, B. 1379. Very 
little of this plant seen. 


600 


GERANIACEAE. 
*ELrodium Botrys, Bert. O. 1612. 
E. cygnorum, Nees. H. 1526, T. 1191, 1744, K. 1821. 


ZYGOPHYLLACEAE. 


Tribulus terrestris, Linne. K. 1842. 

Zygophyllum apiculatum, F. v. M. B. 1377. 

Z. Billardieri, DC., var. ammophilum, J. M. Black. 
OO. 159 Bs TS hb et sie... 

Z. fruticulosum, DC. O. 1233, 1581, 1599\ 7 ee 
1389, W. 1208, K. 1843. . 

Z. wdocarpum, F. v. M. H. 1253, 1509, K. 1844. 

Z. ovatum, Ewart et White. H. 1254, 1562, O. 1580, 

. B. 1353, K. 1845. ‘‘Dis. W.”’ These are new locali- 

ties for this rare plant. 


EUPHORBIACEAE. 
Euphorbia Drummondu, Bois. H. 1613, O. 1572,- 
T1782: 
E. eremophila, Cunn. O. 1592, T. 1783, K. 1822. 
Poranthera microphylla, Brong. B. 1325. ‘‘Dis. W.” 
Adriana Hookeri, (F. v. M.) Muell. Arg. O. 1305. 


STACKHOUSIACEAE. 

Stackhousia muricata, Lindl. O. 1594, B. 1355. There 
is a doubt about this identification, as S. vwminea, 
Smith, apparently only differs in the corolla lobes 
being acute and not obtuse. ‘“‘Dis. W.” 


SAPINDACEAE. 
Heterodendron oleaefoluum, Desf. O. 1590, T. 1787. 
Dodonaea attenuata, Cunn. O. 1631, B. 1224, 1388. 
D. microzyga, F. v. M. OQ. 1281, 1584, B. 


MALVACEAE. 
Sida calyxhymema, J. Gay. T. 1728. i 
S. corrugata, Lindl., var. orbicularis, Benth. H. 1566, — 
1261 On sora sds, ToL Tob, .t 
S. corrugata, Lindl., var. ovata, Benth. W. 1398, 
T. 17195 3738: 
Sida intricata, F. v. M. K. 1837. 
S. wrgata, Hook. T. 1796. 
Abutilon Mitcheliu, Benth. ._T. 1759. ‘Dis. W.” 
A. otocargum, Iw MM. OL 16220" “Dis. Woe 
A. oxycarpum, F.v-M. T. 1727, K. 1800. “Dis. Wee 
Lavatera plebera, Sims. H. 1537, O. 1252, 1295. 
*Malva parviflora, L. T. 1797. 


601 


FRANKENIACEAE. 


_Frankenia pauciflora, DC. H.'1510. “‘Dis. W.”’ 
F. serpyllifolia, Lindl. T. 1716, Ke eeok. 


ee ekouae. 


Pimelea maderneethale, fers O: Lois: B. ied, 'T. 1792. 
Eesupier, Woy, Mo O.:1619, B. 1349. 


MYRTACEAE. 


EUCALYPTUS SPECIES IDENTIFIED BY Mr. J. H. MaiIbDeEn, 


Ps Oso, etc. 

‘E. gracilis, F.v. M. O. 1477-8. (‘‘With large fruits.” — 
ee Mo) bs. W’ 

E. werassata, Labill.(?). I. 1340. (‘‘Perhaps this 
species and close to the tyne.’’—J. H. M.) 

B.-olcosa, KF. vy. M.*’O. 1270, 1473, 1483, L. 1481-2, 
B. 1337, 1346, 1361, 1372. No. 1372. (‘‘This seems 
a very interesting form.” —J. H. M.) Remarkable 
for its prostrate trunks, horizontal branches, and 
narrow grey glaucous leaves Cpl xii te Vay 

E. pyriformis, Turcez. O. 1310, with large fruits 5°6 cm. 
across, and O. 1484, with smaller fruits’ 3°7 cm. 
across. ) 

E. transcontinentalis, J. H. Maiden. O. 1288, 1292, 
1371, 1373, 1475-6, B. 1344. 

i. uncinata, Turez. Gee Ob £2997) >(“ Probablyd torm,”’ 
5 ae M.) 

H., sp. 1. 1480. Mr. Maiden advises that he is making 
a new species of this plant. 


Melaleuca hakeoides, F. v. M. I. 1245. ‘“‘Dis. W.” 


i. parvifiora, Lindl. O. 1304. 

Thryptomene Elliottw, F. v. M. B. 1218, 1312. 

Leptospermum laevigatum, F. v. M., var. minus, F. v. M. 
O. 1278. 

UMBELLIFERAE. 

Uldima mercurialis, J. M. Black. O. 1267. Mr. Black 
is describing this new genus in these Transactions 
this year. I only saw this plant at Ooldea, after 
which it is named. Remarkable for the horizontal 
barbed wings to the fruit. | 

Didiscus glaucifolius, F. v. M. B. 1314, 1347. ‘‘Dis. 
ANE : 

Daucus glochidiatus, (Labill.) Fisch. H. 1560, O. 1610, 
K. 1817. (D. brachiatus, Sieb.) 


PRIMULACEAE. 


*Anagallis arvensis, L. O. 1600, B. 1356. 


602 


ASCLEPIADACEAE. 
Marsdena Leichardtiana, F.v. M. O.1601. ‘‘Dis. W.”’ 


CONVOLVULACEAE. 
1 ba = Fe 3 = 
Convolvulus erubescens, Sims. H. 1528. 


BoRRAGINACEAE. 
Lappula concava, F. vy. M. O. 1307, T. 1789. 
*Tithospermum arvense, L. O. 1632. 
*ERchium plantagineum, L. B. 1378. 


LABIATAE. 

Westringia Dampieri, R. Br., var. rigida, J. M. Black. — 
O. 1630, B. 1375: 

SOLANACEAE. 

Solanum coactiliferum, J. M. Black. I. 1248, B. 1231, 
1327. 

S. ellipticum, R. Br. O. 1624, T. 1794, K. 1838. 

S. orbiculatum, Dunal. B. 1359. 

Anthotroche truwcata, EK. H. Ising. O. 1297, B. 1374. 
This new species is most rare; only two plants of it 
were seen. For description see p. 605 and pls. xxxviil. 
and xxxix., dig: ;1. 

Nicotiana suaveolens, Lehm. H. 1559, O. 1586a, B. 1322. 

Lycium australe, F. v. M. T. 1791. 

Duboisia Hopwoodu, F. v. M. B. 1343. 


MYOPORACEAE. 


Myoporum platycarpum, R. Br. W. 1215, O. 

Kremophia alternfolia, R. Br. O. 1242, 1530, 1533a, 
1579, B: 1710, T. 1402, K..1820. NesabSG>iea8 
shrub 2 metres in height and was remarkable for | 
the variation in the colour of the flowers. On the © 
same bush some flowers were wholly pink or red- 
dish, others were partly pink and partly white, while 
others were all white. 

EL. Duttonu, F. v. M. T. 1406, 1779. 

E. Gabsomi, F v2 M. O: 1265, 1308; ‘Disa | 

EL. glabra, (R. Br.) Ostenf. B. 1321, 1366, 1704, 17113 F 
T. 1415. (H. Brown, F. v. M., is a synonym.) | 

FE. latifolia, Fi v.iM. O.°1587,°T. 1741, Wee } 

E. Latrobei, F. v. M. H. 1291,°°O. 1234a, T. 178GRRe 
K. 1826. | 

EL. Latrobei, F. v. M., var. Tietkensii, Tate.. O. 1234; 
1243. 

E. maculata, F. v. M. H. 1251, 1540, O. 1621: 


} 


603 


E.. Paislem, Bowe Me T) 1412. 
HE. rotundifolia, F. v. M. T. 1403. 
Bs scoparia, ho seo 4B) 1336, TT a78r: 


PLANTAGINACEAE. 
Plantago varia, R: Br!) H. 1522, T. 1197. 


RUBIACEAE. 
Pomaz umbellata, Sol. O. 1586. 


CUCURBITACEAE. 
*Cucumis myriocarpus, Naud. B. 1392. 


CAMPANULACEAE. 
Wahlenbergia gracilis, DC. B. 1320. 


GOODENIACEAE. 


Vellewa paradoxa, R. Br. O. 1615. The calyx consists 
of one ovate toothed sepal 10 mm. long and four 
shorter lanceolate entire sepals 7-8 mm. long. ‘‘Dis. 
Wee? 

Goodema pinnatifida, Schiect. H. 1255, 1551, O. 1616, 
et P3020 TS 1745: 

G. pusiliflora, F. v. M. T. 1405, K. 1827. 

Scaevola spinescens, R. Br. O. 1277, 1597, B. 1219, 1370. 

Calegane perardiana,, Bove MT. 1418; 1736. “Dis. 
W ? 

CoMPOSITAE. 

Olearia Muellert, Benth. O. 1617, B. 1348. (Synonym, 
Aster Muellerr, F. v. M.) 

O. subsyicata, Benth. O. 1266, B. 1381. (Synonym, 
Aster Mitchella, F. v. M.) ‘Dis. W.” 

Vittadinia australis, A. Rich. H. 1525. 

Vo seabra, WC. .O. 1691, B. 1342. “Dis. W.’’ 

Podocoma nana, Ewart et White. H. 1570, K. 1801. 
This rare plant has only been previously recorded 
from Glen Ferdinand, Everard Range, Mount 
Lyndhurst, and Torrens Plain (vide J. M. Black in 
these Transactions, vol. xxxix., 1915, p. 839). 
Dist We? 

Minuria Cunningham, Benth. H. 1573. “‘Dis. W.”’ 

M. leptophyla, DC. H. 1514, O. 1635, K. 1802. 

Calotis cymbacantha, F. v. M. W. 1211, 1397, T. 1199, 
1754. 

C’. erinacea, Steetz. O. 1693. 

C. hismdula, F. v. M. H. 1523, B. 1391, T. 1770. 

| C. multicaulis, (Turcz.) J. M. Black. T. 1419. 

$2 


504 


C. multicaulis, (Turez.) J. M. Black, nov. var. brevi- 
radiata. Variat lguhs radu brevissimis, disco 
achaenu glabro absque ayice pubescente, cilus alarum 
sursum prominenter lobatarum simplicibus, papp 
aristis sine barbellis reflexis, folus wferioribus 
angustioribus et acutius dentatis. H. 1552. Differs 
in the ligule of the ray flowers being very short, in 
the achenes being almost glabrous except for the 
pubescence at the summit, the hairs on the wing- 
margin simple, the wings prominently lobed at the 
top, and the awns without reflexed barbs, the lower 
leaves narrower and more sharply toothed. 

Brachycome ciliaris, (Labill.) Less. O. 1309, 1583, 
B. 1709). Wi) 14005 Devi 7685) Ks Vedag 

B. ciliaris, (Labill.) Less., var. glandulosa, Benth. — 
OL Wet | .| 

B. Muellert, Sond. W. 1205, 1396, T. 1421. “Dis. Was 

B. pachyptera, Turcz. T. 1747, K. 1815. 

*Centaurea militensis, L. K. 1813. ' 
Cratistylis conocephala, 8. Le Moore. O. 1274, T. 1755. 
Elachanthus pusillus, F. v. M. H. 1527, K. 1818. ; 
Tsoetopsis gramuufolia, Turcz. H. 1524, K. 1825. “Dis. — 
Myriocephalus Stuartu, Benth. O. 1306, T. 1193, 1713. 
Siloxerus brachypappus, (F. v. M.) comb. nov. H. 1529, 

O. 1699. As Srloxerus, Labill., is the earlier name 
(1806) it must replace Angianthus, R. Br. (1810). | 
Mr. J. M. Black drew attention to this in his “‘Flora ~ 
of South Australia’’ (1922), p. 6. ; 

S. pusilius, (Benth.) comb. nov. O. 1240, B. 1323. 

Gnephosis cyathopappa, Benth. T. 1765, K. 1853. © 

G. skirrophora, Benth. H. 1518, 1553, O. 1695, K. 1852. 

Gnaphalodes uliginosum, A. Gray. O. 1290, 1585, 
By. Pab8 ol, Urea, 

Craspedia pleiocephala, F. v. M. W. 1212, T. 119455 
17A2, oS, Kose 16. 

Eriochlamys Behru, Sond. et F. v. M. T. 1731. 

Toxanthus Muellert, Benth. B. 1324. ‘‘Dis. W.” 

Rutidosis helichrysoides, DC. K. 1847. > 

Millotia Kempei, F. v. M., var. Helmsii, F. v. M. eb ¥ 
Tate. O. 1576. | Si 

M. tenwifolia, Cassini. T. 1715. 


oo ae 


Pe & eds 


Ce ee 


Txiolaena leptolepis, Benth. K. 1849, 1850. i 
Podotheca angustifolia, Less. O. 1694. 3 | 


Podolepis acuminata, R. Br. T. 1720. “Dis. W.”’ 

P. canescens, A. Cunn. H. 1558. a 

P. capillaris, (Steetz.) Diels. B. 1220, 1326, W. 1203, | 
T. 1738; 


=. 


i 


605 
Leptorhynchus tenuifolius, F. v. M. O. 1696. ‘‘Dis. 
w 9 


L. tetrachaetus, (Schlect.) J. M. Black, var. penicillatus, 
J. M. Black. K. 1848. 

Helichrysum ambiguum, Turcz. T. 1773, 1734. | ‘Dis. 
We? 

H. ayculatum, DC. O. 1300. 

H. Lawrencella, F. v. M. O. 1582, B. 1223, 1225., 

H. Lawrencella, F. v. M., var. Davenport, Benth. 
i. 1786: 

H. lucidum, Henck. B. 1352. 

H. Mellorianum, J. M. Black. I. 1250. 

Waitzia acuminata, Steetz. O. 1596, B. 1703. 

Helipterum Charsleyae, F.v. M. K. 1804. ‘‘Dis. W.”’ 

. Pitzgibbonn, F. v. M. T. 1409. 

. floribundum, DC. H. 1538, O. 1595, B. 1316, 1363, 

W. 1210, T. 1420, K. 1823. 

. Humboldtianum, (Gaud.) DC. O. 1692, T. 1401. 

wisiessenie, “Why. ME MK T8038: 

. moschatum, Benth. T. 1198, 1410, W. 1399. 

. pterochactum, Benth. T. 1750. 

. pygmaeum, Benth. K. 1828. 

. roseum, (Hook.) Benth., var. patens, (Hwart) J. M. 

Black. W. 1206. 

 stipitatum, Fv. M. K. 1854. “Dis. We? 

. strictum, Benth. H. 1519, K. 1824. 

. tenellum, Turcz..: H. 1568. 

ietremain wee Ne Oy P6383." By. 1850: 

Senecio brachyglossus, F. v. M. H. 1571, O. 1634. 

Hea Gnegorn., Bev. MeO T62T) W209) T1168: 


Ry 


Ree RPRRRRA 


*Cryptostemma calendulaceum, R. Br. O. 1296. 


Cephalipterum Drummondu, A. Gray. H. 1535, T. 1730, 
1735. 


VII. A New Soranacrtous Puant, 
Anthotroche truncata, n. sp. 
Pise xxvii. end, xo. fio. 1. 
Frutex bimetralis milis brevissimis plumosis divaricatis 


dense obtectis, folus oblongis vel ovatis 5-10 mm. longis 
obtusis incanis breviter petiolatis, nervo medio prominente, 
floribus subsessilibus odoratis, calyce tabulato viz 2 mm. longo 
— lobis ejus deltoiders brevissimis, corolla alba extra tomentosa 

3 strus longitudinalibus signatd intus glabra lobis ejus late 
- oblongis patentibus tubum subaequantibus, filamentis basi 
_ dilatatis et pilosis, ovario parce stellato-piloso. 


Ooldea, East-West Railway Line, September 15, 1920, 


_ and Barton, in the same district, September 19, 1920. 


, 606 


A handsome shrub of 2 metres high, hoary. Branches 
round, hoary, with a fine tomentum wearing off in age, divar- 
icate. Leaves 5-10 mm. long and 3-5 mm. wide, broad oblong 
to ovate, sometimes broad at base, entire, obtuse, hoary, with 
a very fine down of plumose hairs, midrib above and below 
and often a few lateral veins, prominent, scattered, or in 
clusters of 2 or 3, petiole very short. Flowers 1 to 3 in the 
leaf. clusters, almost sessile, sweet-smelling. Calyx tubular, 
1°5 mm. long, investiture similar to the leaves; lobes very 
short, obtuse triangular. Corolla white with three fine reddish 
short longitudinal lines inside, tomentose outside, except 
almost smooth near the base, glabrous inside; lobes oblong, 
as long as the tube, spreading. Stamens 5, hardly exsert; 
filaments dilated and pilose at the base. Ovary with a few 
stellate hairs; ovules 2 to 3 in each cell, only one appears to 
develop and is finely tuberculate. 3 

The new species is nearest to A. Blachu, F. v. M., but 
differs from this and all other species of this genus in the 
truncate calyx, the tomentose clothing, and rotate corolla. 


DESCRIPTION OF PLATES. 


Pratt XXXVITI. 
Anthotroche truncata, n. sp. 1, Flower, viewed near the top; 
2, flower, side view, showing calyx; 3, stamen, showing dilated 


pilose filament. 
PrateE XXXIX. 


Fig. 1. A new solanaceous plant (Anthotroche truncata, 
n. sp.) growing on a sandhill at Ooldea showing habit. 

Fig. 2. Vegetation on a sand ridge at Ooldea showing 
Eucalyptus transcontinentalis, Eremophila alternifolia, Olearia 
Muellerit, Triodia irritans, and Westringia Dampier. 


Puatse XL. 

Fig. 1. Nullarbor Plain at Hughes showing the open forma- 
tion of bluebush (Kochia sedifolia) and saltbush (Atriplex 
wesicarium ). 

Fig. 2. At Ooldea Soak showing a carpet of Myriocephalus 
Stuarti with Leptospermum laevigatum, var. minus, on the right. 

Pruate XLI. 
Fig. 1. Eucalyptus oleosa at Barton with prostrate trunks. 


Fig. 2. Barton from a sand ridge with Thryptomene Elliotti 
in the foreground, Casuarina lepidophloia and Eucalyptus below. 


Prate XLIT. 

Fig. 1. Looking north of Tarcoola showing the stony, un- 
dulating nature of the country. Acacia tarculensis and Trichinium 
meanum in the foreground. 

Fig. 2. On the flats at Kingoonya. Minuria leptophylla in 
the foreground with trees of Acacia Loderit and shrubs of Kochia 
sedifolia in the middle distance. 

The photographs were taken by myself. 


607 


MISCELLANEA. 


Note on Diastoma melanioides, Reeve (Mesalia). 
By Siz Josepn Verco, M.D. (Lond.), F.R.C.S. (Eng.). 


DIASTOMA MELANIOIDES, Reeve. 


Mesalia melanioides, Reeve, Conch. Icon., vol. v., pl. i., f. 3. 
ee (?) KE. A. Smith, Ann. and Mag. Nat. Hist. Ser. 8, vol. 
-AGIS, p. 310. 


" Mesalia extlhis, Sowerby, aa and Mag. Nat. Hist., vol. xii., 
py. 2360, ply 111. , fig. 9, W. Austr. 


This shell was dredged by me in 1895 in 15 Eabh one off 
_ Thistle Island, at the entrance to Spencer Gulf, with two 
smaller examples, and measured 42 mm. in length and 
11°25 mm. in breadth. A dead shell was found on the Thistle 
Island beach. Off the Banks Group, in Spencer Gulf, in 
12 fathoms, one small fresh example was dredged and one of 
medium size dead. Later four specimens were taken on St. 
_ Francis Island beach, the largest of which, in perfect con- 
_ dition, was 41 mm. long and 12°5 mm. wide. In Petrel Bay, 
on the north of the island, in 15-20 fathoms, five very small 
dead specimens were dredged, and in 6 fathoms three tips. 
In 1911, at Esperance Bay, on the south coast of Western 
Australia, six full-grown beach specimens were obtained 
measuring up to 42°25 mm. long by 12°75 mm. wide. Shortly 
afterwards one of the latter was given to Mr. G. B. Sowerby 
when on a visit to Australia, as an example of Jf. melanioides, 
Rve., from Esperance, and a little while after this a reprint 
was received from him containing the publication of his 
M. exilis. When reminded of the circumstances under which 
he obtained it, he explained that he had failed to make a note 
of them at the time and they had slipped his memory, and. 
without doubt his name was a synonym of Jf. melanioides, 
Rve. Its type locality is Esperance Bay. 

The whorls in some examples are nearly flat and sloping, 
in others slightly convex ; with a finely canaliculate suture, and 
_ with a shallow spiral groove about one-fifth the width of the 
- whorl below the suture, which consequently seems somewhat 
marginal or adpressed. The numerous broad rounded axial 
costae are very valid in the upper whorls, where many of them 
_ are variceal, and these may form in some examples three vertical 

_ lines of varices, each just in front of that above; in other 


608 


examples they are quite irregular. The varices disappear in 
the later whorls, and the axial costae also gradually fade out. 
The spiral lirae (with two to five intervening striae), about 
six in the spire whorls and twice as many in the body whorl, 
retain their validity. The thickly-glazed inner lip gives the 
impression that the callus of the posterior half has been first 
laid down over a circular area, and the anterior half laid 
down upon this over an area with a shorter radius, so that 
the edges of the areas meet each other at a wide angle, and 
the edge of the anterior circular area is continued into the 
aperture as a raised curved plait or carination. Its lower edge 
curves round anteriorly, and forms with the basal lip a 
shallow wide sinus with a slightly everted edge. The proto- 
conch consists of two smooth convex homostrophe whorls. The © 
ornament is composed of squarish light-chestnut spots imme- 
diately below the suture, with smaller spots more or less 
distantly articulating the lirae, and sometimes also so disposed 
as to form curved axial narrow flames of dots. 

It is very closely allied to the fossil Diastoma provisi, 
Tate, Journ. and Proc. Roy. Soc. N.S. Wales, vol. xxvii., 
1893, p. 177, Miocene and Older Pliocene (now recognized 
as Older and Newer Pliocene). Tate diagnoses between the 
two. He also shifts both species from the genus Mesalia 
to Diastoma, Deshayes. He writes, ‘‘Cossman, to whom the 
fossil was sent under the above name” (Mesalia provisi), 
‘“Gnforms me that it is a Diastoma; from him I have received 
examples of several species of Diastoma and Mesalia from the 
Parisian Eocene. This material permits me to affirm that 
M. provisi, Mihi, and Jf. melanoides, Rve., are congeneric 
with D. costellatum, Lamarck; whilst MJesalia sulcata, 
Lamarck (non sulcata, Gray=brevialis, Lamarck), is of a 
totally different type. Diastoma simulates Mesaha, but the 
latter has a sinuated outer lip, whilst the spiral carination 
of the columella of Diastoma is quite a different feature from 
the slight twist of the columella-margin of Mesalia; more- 
over, Diastoma is more or less variced. Mesalia belongs to the 
‘ Turritellidae; Diastoma, which has been located in at least 
two families, finds a resting place in Cerithiidae, it may be 
viewed as a Melania-like Cerithium.”’ 


E. A. Smith, in his review of the Genus Wesalia, loc. 
cit., swpra, does not refer to Tate’s transfer of J/. melanioides, 
Rve., to the Genus Diastoma, which, how ores merits notice 
and acceptation or refutation. 


Evening Meeting, September 14, 1922. 


- pag 


609 


An Introduced Land Snail, Helicella ventricosa, 
Draparnaud. 
Sir Joseph Verco showed a number of small snails col- 


lected in a garden at Woodville, at the end of last month. 
_ They were first noticed about five or six years ago in a bed 


of petunias, which they completely destroyed by ringbarking 


_ the stems rather than by consuming the leaves. They belong 


to the same species and are of the same size as some snails 
sent to the Adelaide Museum from Mount Gambier, which 
were identified as Helicella (Cochlicella) ventricosa, Drapar- 
naud. Their habitat is the south of Europe and the north of 
Africa, the Canary Islands, and the Azores. They are found 


_also in Bermuda as an introduction. They have evidently 


been brought by some means into South Australia, where they 
appear to be now widespread and numerous. A note of their 


_ appearance as a novelty near Mount Gambier is found in the 
last issue of the Records of the South Australian Museum, 


vol. 11., No. 2, April 3, 1922. 
Jos: C. Varco: 
Evening Meeting, May 11, 1922. 


610 


ABSTRACT ‘OF “PROCEED 


Royal Society of South Australia 
(Incorporated) 


FOR THE YEAR NOVEMBER 1, 1921, To OcToBER 31, 1922. 


ORDINARY MEETING, NovEMBER 10, 1921. 
THE PRESIDENT (R. S. Rogers, M.A., M.D.) in the chair. 


THE PRESIDENT referred to the approaching centenary of 
the Royal Society of New South Wales, and it was resolyed— 
‘That a suitable letter of congratulation be forwarded to that 
Society.” 

ELEctTions.—Owen M. Moulden, M.B., B.S.; Melville 
Birks, M.B., B.8., L.R.C.P., F.R.C.S.; Professor T. Harvey 
Johnston, M.A., D.Sc.; and Oscar W. Tiegs, M.Sce., as 
Fellows. 

PapEers.—‘‘The -Pathological Morphology of Cintractia 
spimficis,’’ by Prof. T. G. B. Ossorn, D.Sc.; ‘Occurrence 
of Remains of Small Crustacea in the Proterozoic(?) or Lower 
Cambrian(?) Rocks of Reynella, near Adelaide,’ by Prof. 
Sirk Eperworts Davin, D.Sc., F.R.S., etc. ; ““A New Species 
of Lycosa for South Australia,” by R. H. Pulleine, M.B. 

Exuisits.—Mr. L. Keita Warp showed lantern slides 
of Typical Views of the Eucla Basin and Nullarbor Plain, 
with Maps descriptive of the topography, geology, rainfall, 
and artesian water supply of the district. Mr. A. M. Lza 
exhibited the three known blind beetles of South Australia, 
Illaphanus stephensi (Carabidae), Rodwayia minuta, Lea 
(Tricopterygidae), and Halorhynchus caecus (Curculionidae). 
Capt. S. A. WuitEe showed botanical specimens from the 
North-western District of (New South Wales. Sir DovuGuas 
Mawson showed calcareous deposits from a series of caves im 
the limestone near Reynella. 


OrpInARY MEETING, ApRiL 13, 1922. 
THE PRESIDENT (R. 8S. Rogers, M.A., M.D.) in the chair. 
THE PRESIDENT welcomed as a visitor Dr. McGillivray, 


President of the Broken Hill Ornithological Society. He also 
announced with deep regret the death of Mr. F. RB. Zietz, 


611 


who had joined the staff of the South Australian Museum 


more than thirty years ago, and was at the time of his death 
ornithologist to that institution. He was elected a Fellow of 


e 
z 


Sas 


>. 


a 


ety 


_ to travel in the Board’s s.s. ‘Victory. 


this. Society in 1912, and had contributed important papers 
on the Wild Hybrids of Australian Ducks, and Australian 
Lacertilia: he had also taken an active part in the discus- 
sions, and been responsible for many interesting exhibits at 
the meetings. 

Evections.—J. Sutton and Thos. Draper Campbell, 
B.D.S., were elected Fellows. 


Papers.—‘‘Notes on Australian Polyplacophora, with 
descriptions of three New Species and two New Varieties,’’ 
by Epwin Asusy, F.L.S., M.B.O.U.; ‘‘A New Isopod from 
Central Australia belonging to the Phreatoricidae,”’ by Cuas. 
Cuitton, D.Sc., C.M.Z.S. (communicated by Prof. F. Wood 
Jones, M.R.C.S., D.Sc., etc.); ‘‘The Flora and Fauna of 
Nuyt’s Archipelago and the Investigator Group, No. 1— 


_Amphipoda and Isopoda,”’ by Cuas. Cuitton, D.S¢., 


C.M.Z.S. (communicated by Prof. F. Wood Jones, M.R.C.S., 
D.Sc., etc.) ; ‘‘The External Characters of Pouch Embryos of 
Marsupials, No. 3—J/soodon Barrowensis,’’ by Prof. F. Woop 
Jones, M.R.C.S., D.Sc., etc. 


REso_veD—‘‘That the types and co-types collected by 


_ Prof. F. Wocd Jones during his exploration of Nuyt’s 


Archipelago be presented to the South Australian Museum.” 


RESOLVED—‘‘That a letter be sent to the Chairman of 
the S.A. Harbours Board expressing appreciation of the 
courtesy extended to the Professor by affording him facilities 


Exuisits.—Mr. A. M. Lea exhibited a collection of 
bones taken from the pellets of the common owl, representing 


a year’s food for one of these useful birds. It included bones 
of 1,407 mice, 143 rats, 5 young rabbits, 375 sparrows, 23 


_ starlings, and a few other birds, frogs, and bats; also some 


insect remains. Some of the bones showed considerable sponge- 
like swellings, indicating serious disease. He also showed a 
collection of insects from North-west Australia, presented by 
Dr. Morgan. Also a root from Mr. L. Harnett, taken from 
the ground under an Adelaide building erected 62 years ago, 
and still perfectly sound. Prof. F. Wood Jones exhibited 


_ three maxillae of Thylacoleo and portions of maxillae and 
_ mandibles of Thylacinus which he had found in Buckalowie 
_ Cave, No. 2, near Carrieton. Capt. S. A. Wuite showed a 


large sheet of mycelium of a remarkable fungus resembling 
chamois leather, found between the layers of wood of a giant 
Eucalyptus rostrata felled at the Reedbeds, near Adelaide. 


612 


OrpDINARY MEETING, May 11, 1922. 


Tue Presipent (R. 8S. Rogers, M.A., M.D.) in the chair. 

The Hon. Secretary stated that the Council had asked 
Prof. Sir William H. Bragg to represent the Society at the 
150th Anniversary of L’Académie Royale de Belgique, and 
read a reply regretting his inability to be present on ee) 
occasion. 

The Hon. Srcretrary read a letter from the Natiostal 
League for the Protection of Natural Monuments, Florence, 
asking the co-operation of the Fellows in the provision of 
illustrations of native animals for zoological handbooks to 


be issued by the League. 


Papers.—‘‘A Geological Traverse of the Flinders Range 
from the Parachilna Gorge to the Lake Frome Plains,’’ by 
Prof. WALTER Howcuin, F.G.S.; and ‘‘The Parasites of Aus- 
tralian Birds,’’ by Prof. J. Burton CLeLtanp, M.D. In the 
absence of the author, and also, through illness, of Prof. F. 
Wood Jones, who was to have introduced it, the latter paper 
was taken as read. 

Exuisits.—Sir JoserH C. Verco showed a number of 
small snails (Helicella ventricosa, Draparnaud) (vide Mis- 
cellanea). Mr. A. M. Lea exhibited a drawer of rove beetles 
(Staphylinidae), several of which have remarkable combs on 
the middle leg. One species lives on bush rats, another on. 
the flying fox, and others in nests of ants. Mr. W. J. KimBer 
showed large fossil sharks’ teeth from Port Willunga cliffs; 
polyzoic limestone and a cast of a large cowrie shell from Point 
Turton; and Truncatilla scalarina, associated with large 
deposits of sepia bones in raised beach at Minlacowie. 


OrpDINARY MEETING, JUNE 8, 1922. 
THE Presipent (R. S. Rogers, M.A., M.D.) in the chair. 


Nominations.—C. T. Madigan, B.A., B.Se.; Guy A. 
Lendon, M.B., B.S., M.R.C.P.; and Alan H. Lendon were 
nominated as Fellows. . 

Paprers.—‘‘The Tertiary Brown Coal-bearing Beds of 
Moorlands,” by Prof. Str Doucitas Mawson, D.8c., B.E., 
and FREDERICK CHAPMAN; ‘‘External Characters of Pouch 
Embryos of Marsupials, No. 4—Pseudochirops dahli,”’ by Prot. 
F. Woop Jones, M.R.C.S., D.Sc., etc.; and ‘‘A New Species 
of Puecima ” by C. F. JOHNCOCK, Corr. Mem. (conte 
by Prof. W. Howchin, F.G.S.). 

Exuisits.—Prof. W. Howcuin exhibited several highly- 
glaciated erratics obtained during his late expedition into | 
Central Australia in company with Prof. Sir Edgeworth | 
David under a grant from the Australian Association for the 


i 613 


Advancement of Science. The specimens were obtained from 
the tillite exposed at Yellow Cliff on the Finke River, and 
from a new locality in the Crown Point Hill Range in the 
same neighbourhood. Mr. A. M. Lea exhibited a common 
Queensland scorpion (Hormurus caudicula) obtained alive in 
Adelaide by Mr. A. Bottcher in some bananas from 
Queensland: 
ORDINARY MEETING, JULY 13, 1922. 


THE PresipEnT (R. S. Rogers, M.A., M.D.) in the chair. 
Nominations.—Geofirey Samuel, B.Sc.; Wiliam Ham, 
ipo. and nh, i. T. Grant, M.B., B.57,’ M:R-C.P., as 
Fellows. 
4 Evections.—C. T. Madigan, B.A., B.Sc.; G. A. Lendon, 
» M.B., B.S., M.R.C.P.; and A. H. Lendon, as Fellows. 
Papers.—‘ ‘Contributions to the Orchidology of Aus- 
tralia and New Zealand,’ by R. 8S. Rocers, M.A., M.D.; 
| ‘‘The Physiography of the Meadows Valley, Mount Lofty 
'~ Ranges,”’ by E. O. Teatze, D.Sc. (communicated by Prof. 
_ Walter Howchin, F.G.S.) 
Exuisits.—Mr. A. M. Lea exhibited a drawer of insects 
_ showing remarkable differences in the sexes. Prof. J. B. 
_ CLELAND exhibited a specimen of the rare puffball Mytremyces 
| _ fuscus, Bert. This is a new record for South Australia, and 
__ was found on the shady side of a road cutting, at Mount 
. Lofty, on July 1. The exhibitor has only personally met 
_ with this fungus once before, on a similar shady bank on 
the Cambewarra Mountains, near Noura, N.S.W. The plant 
has an erect dirt-coloured fenestrated stem, capped by a 
rounded receptacle showing projecting whitish (sometimes 
vermilion) teeth. In an early stage the receptacle is covered 
_ with a little cap, one of which was also exhibited, having 
_ been thrust off on to the ground. Dr. E. Anecas JOHNSON 
_ showed two specimens of Umio from River Onkaparinga. 
Prof. F. Woop Jones showed two adult specimens of 
Myrmecobius, probably distinct races, one from Western Aus- 
tralia and one from South Australia. Mr. E. R. Waite 
_ showed a model, one-tenth natural size, of Camarasaurus, and 
_ a fossil femur of the same Dinosaur, found at Wyoming, 
U.S.A. The model was prepared under the direction of Prof. 
_ Henry Fairfield Osborne, Director of the American Museum 
| of Natural History. Mr. R. L. Jacx showed a Bootes 
_ model of Iron Knob and its vicinity. 


_ ORDINARY MEETING, Aveust 10, 1922. 


THE PresIDENT (R. S. Rogers, M.A., M.D.) in the chair. 
Exvections.—R. L. T. Grant, M.B., B.S., M.R.C.P.; 
| William Ham, F.R.E.S.; and Geoffrey Samuel, B.Sc., as 
: Fellows. 


614 


Nominations.—Miss E. D. Nobes, B.Sc.; Herbert M. 
Hale; and Albert Geo. Charles, as Fellows. 

THe PRESIDENT reported that at the invitation of the 
Queensland Branch of the Royal Geographical Society the 
Council had appointed Prof. F. Wood Jones to represent 
them upon a joint committee to consider the suggested ex- 
ploration of the Great Barrier Reef, and Prof. Sir Douglas 
Mawson to represent the Society on the Council of the Aus- 
tralasian Association for the Advancement of Science at its 
meeting in Wellington, N.Z. Also that a letter had been 
received from Queensland asking for co-operation in urging 
the Government to take steps to prevent the extinction of 
the Ceratodus, and that the Council had endorsed the sug- — 
gested action. 

PaPerRs.—‘‘Some New Records of Fungi from South 
Australia, Part II., together with a description of a New 
Species of Puccima,’’ by Prof. T. G. Ossporn, D.Se., and 
GEOFFREY SAMUEL, B.Sc.; ‘““An Investigation of the Essential — 
Oil obtained from Hucalyptus cneorifolia, DC.,”’ by PHiLip 
A. Berry, B.Sc. (communicated by Prof. E. C. Rennie, 
D.Sc.); ‘‘The Flora and Fauna of Nuyt’s Archipelago and 
the Investigator Group, Part 2—The Monodelphian 
Mammals,” by Prof. F. Woop Jones, M.R.C.S., D.Sce., 
etc.; “‘The Flora and Fauna of Nuyt’s Archipelago and 
the Investigator Group, Part 3—A Sketch of the Ecology of 
Franklin Island,’’ by Prof. T. G. B. Ossorn, D.Sc. 


ORDINARY MEETING, SEPTEMBER 14, 1922. 


Tue Presipent (R. S. Rogers, M.A., M.D.) in the chair. 

Exections.—Miss E. D. Nobes, B.Sc.; H. M. Hale; 
and A. G. Charles, as Fellows. 

Papers.—‘‘On the Striation of Voluntary Muscle Fibres 
in Double Spirals,’ by O. W. Trees, M.Sc; “‘The Flora 
and Fauna of Nuyt’s Archipelago and the Investigator 
Group, Part 4—Coleoptera,’’? by Antoun M. Lua, F.E.S.; 
‘‘Australian Lepidoptera of the Tribe Geometrites,” by A. 
JEFFERIS TurRNER, M.D., F.E.S.; ‘‘Australian Coleoptera, 
Part III.,. by Ausert H. Exston, F.E.S.; ‘“‘Cylindro- 
Conical Stones from the Darling River and Cooper Creek,’’ 
by R. H. Putieine, M.B. : 

Exuisits.—Sir JosePpH VERCcO showed some shells (vide 
Miscellanea) ; also 27 almonds in their shells taken from the 
crop of a game rooster which was found in convulsions. The 
crop was opened by the owner, a gardener, the almonds 
removed, and the wound sewn up, the fowl being found quite 
lively the next day. Mr. A. E. Epquist showed two samples 
of Loranthus exocarpus, one grown on an orange tree and one 


615 


on a tugasaste. Capt. S. A. Ware exhibited three birds 
taken during his recent transcontinental trip:—Barnardius — 
macgulivrayt (Cloncurry parrot) and Ayprosmiectus cry- 
thopturus (Red-winged parrot), from North-west Queens- 
land, and Barnardius zonarius myrtae (Mrs. Morgan’s parrot), 
from the Northern Territory. Mr. A. M. Lea showed larvae 
of cockchafers from Nantawarra, where they were stated to be 
destroying from 50 to 75 per cent. of the crop on one farm 
by eating the roots. 
AnnuaL Meetinc, Octoser 19, 1922. 

THE PRESIDENT (R. 8. Rogers, M.A., M.D.) in the chair. 

The AnnuaL Report and FINANCIAL STATEMENT were 
read and adopted. ) 

The Field Naturalists’ AnNnuaL Report was read and 


adopted. 


PRESIDENT’S ADDRESS.—The retiring President delivered 
an address, the subject of which was ‘‘A History of the Royal 
Society of South Australia, particularly in its relation to 
other Institutions in the State.”’ 


PRESIDENTIAL ADDRESS. 
By R. S. Rocers, M.A., M.D. 


A History of the Society, particularly in its Relation 
to Other Institutions in the State. 


An annual Presidential address has by no means been an 
established rule in this Society, and during the last forty-six 
years there have been no less than twenty-nine occasions on 
which :t was cmitted. 

While it would seem unnecessary that your President 
should address you as a matter of duty every year, there would 
appear to be good reasons why the observance should not be 
allowed to fall wholly into abeyance. It is obviously a wise 
thing to make a halt in our proceedings now and then, in 
order that a retrospect may be made. Facts and events are 
but too easily forgotten, and occasional opportunity should 
be afforded to record them in their historical sequence. 

As a Society we are no longer young; we have already 
reached our three score and ten, and are hastening towards 
that century which many of you will doubtless celebrate. For 
this reason I desire to direct your attention to some of the more 
salient points in our history, particularly in its relation to 
other institutions in the State. 


1. PRELIMINARY. 


_ It is hardly necessary to remind you that we are the 
immediate offspring of the Adelaide Philosophical Society, 


616 


which became transmuted into the Royal Society by the 
_ simple device of changing its name and some of its laws, ‘but 
which has otherwise led a continuous and unbroken existence 
for seventy years. There were, however, earlier organizations, 
more or less related to our predecessor in their objects and 
personnel, which may in a sense be regarded as end-products 
of their period. These were all ephemeral. They appeared 
upon their little stage, fulfilled in varying degree a useful 
purpose, then vanished into the limbo of history. They are 
even now, after a comparatively brief lapse of time, a little 
difficult to unearth; and when the preliminary spade-work is 
done, their aliases and their fusions and their recrudescences 
make identification in some instances rather perplexing. 


2. EARLY ORGANIZATIONS AND PRECURSORS OF THE ADELAIDE 
PHILOSOPHICAL SOCIETY. 


Not the least important of these precursors of the Philo- 
sophical Society was the South Australian Literary and 
Scientific Association, founded in London in August, 1834, 
just a fortnight after the Bill for the Establishment of the 
Colony had received Royal assent. 

Owing to the good offices of Mr. Thomas Gill, we have in 
the Archives of the Public Library the first minute book of this 
Association. Among the signatories to the form of obligation 
the following names are of special interest :—Dr. John Brown, 
Thomas Gilbert, Robert Gouger, R. D. Hanson, G. S. King- 
ston, Osmond Gilles, Daniel Wakefield, John Morphett, J. 
W. Childers, Raikes Currie, C. G. Everard, R. Torrens, J. 
Hindmarsh, Chas. Mann, B. T. Finniss, and others. Some 
of these men subsequently became active members of the 
Philosophical Society. 

The objects of the Association were: “The Cultivation 
and diffusion of useful knowledge throughout the Colony” ; 
and as a means to this end, one of their earliest acts was the 
acquisition of a small but excellent library, containing books 
of travel and reference, likely to be of special service to a 
young community. 

Sir Charles 8. Napier, the hero of Scinde, was elected as 
President, and for more than a year numerous meetings were 


held at short intervals in London. Some of these were ofa — 


conversational character ; at others addresses were delivered on 
scientific subjects, such as the geology and anthropology of 
Australia. In December, 1835, just prior to embarkation for 
the new Province, a committee was appointed for the ensuing 
year and the records abruptly ceased. The library was packed 
in the same chest as the Royal Charter, and ultimately arrived 


i? 


— 


=o 


617 


in Adelaide, after various misadventures, in a somewhat 
damaged condition. 


According to a statement made by Charles Mann, the 
pressure of employment, incident upon the earlier stages of 
immigration, prevented any further meetings of the Associa- 
tion in the new Colony. 


. It so happened that in 1838, there became established at 
the “Rooms of the South Australian School Society,’’ in this 
city, “The Adelaide Mechanics’ Institution,’ under the 
presidency of James Hurtle Fisher, at that time Resident 
Commissioner and Registrar. The aims of this body were the 
delivery of evening lectures, together with the control of a 
reading room and circulating library of some 300 books. 
Unfortunately it did not receive the support it had anticipated, 
and in less than a year it was in dire difficulties, unable to 
meet its obligations, and consequently in danger of having 
its books sold by public auction. At this critical period of its 
history, the trustees of the South Australian Literary and 
Scientific Association came to its rescue with an offer of 
amalgamation. The offer was accepted. The latter associa- 
tion was dissolved and its library was handed over in trust to 
the new body, which now bore the cumbrous title of “The 
Adelaide Literary and Scientific Association and Mechanics’ 
Institute.’’ 

But these were not healthy days for the survival of such 
societies, and in turn the new venture faded away, rather than 
dissolved. It finally became extinct in 1844. By some means, 
the books which had been brought from England, were 
deposited with Mr. Da Costa to cover a debt of £20, and were 
still in his hands when yet another organization appeared. 


This was the “South Australian Subscription Library,’ 
which was founded in the year just mentioned. Charles Mann, 
in his evidence before the General Committee of the Adelaide 
Library and Mechanics’ Institute some years later, says that 
he and some of his co-trustees of the early London association 
paid the debt due to Da Costa and presented the books to the 
South Australian Subscription Library, ‘‘of which they 
formed the nucleus.’’ In order that they might not be subject 
to any risk consequent upon a dissolution of the society, it 
was stipulated that in such an event they should become 
public property and be vested in three of the principal officers 
of the Colony. 

The Adelaide Subscription Library was modelled on the 
lines of some of the best English institutions. Its subscription 
was high and its membership exclusive. It was unsuitable for 


_ a young colony where the population was small and the number 


618 


of leisured intellectual people very few. As might have been 
expected, it soon began to decline. 


In 1847 a rival arose, with a freer and more democratic 
constitution. The latter was known as the Mechanics’ Insti- 
tute, and appears to have had no connection with the former 
society which bore the same name. 


The rivalry which existed between these two bodies. was 
not of a healthy character, and did not tend to promote the 
success of either. It meant the support by a not very wealthy 
community of two institutions instead of one, and it soon 
became evident, that unless they could in some way combine 
their efforts and resources, both were doomed. 


The Mechanics’ Institute was the first to make overtures 
for a coalition, but these were coldly received by its rival. 
When, however, these overtures were backed by a promise of 
two substantial donations of £100 each from wealthy citizens, 
the proposal was more favourably considered, and, after much 
parleying, a junction was effected. 


Thus was born, in 1848, “The South Australian ‘Liver 
and Mechanics’ Institute. ” Reorganization, however, did 
not prove a panacea for the troubles which had so constantly 
dogged the steps of these various institutions. The amal- 
gamated society showed but short-lived virility. It shifted 
from Peacock’s Buildings, in Hindley Street, to a more central 
position in Green’s Exchange, a site now occupied by the 
Australian Mutual Provident Society. In a very few years, 
owing to mismanagement and other causes, it was in financial 
difficulties. Contrary to expectations, however, it did not 
expire from inanition, as its predecessors had done, but 
suddenly gave birth to a lusty infant, which was to become 
chief partner in a body corporate, with the Philosophical 
Society as a junior member. ‘This influential partnership 
lasted for a quarter of a century, when it was dissolved by the 
Public Library Act of 1884. 


It is hardly necessary to inform you that this infant was 
the South Australian Institute. 


3. Tur ADELAIDE PHILOSOPHICAL SOCIETY. 


(a) Historical Records. 


The early struggles and activities of the Adelaide Philo- 
sophical Society are recorded in its Annual Reports, in the 
newspapers of the day, and in certain documents recently 
transferred by our Society to the Archives Department of the 
Public Library. 


: 619 


These documents comprise : — 


A.—The first minute book of the Adelaide Philosophical 
Society (1853). 


B.—Papers read and deposited with the Society from 
1853. 

C.—Correspondence relating to a proposed Exploring 
Expedition from Fowler’s Bay into the Interior 
(1855). 

D.—Miscellaneous papers and newspaper cuttings. 


E.—Papers relating to its incorporation with the 
S.A. Institute (1856-71). 

F.—Papers relating to the proposed division of the 
Society into Sections (1859). 


G.—Correspondence between the Adelaide Philosophical 

Society and the S.A. Institute on the one hand; 

and between the Royal Society and Public Library 
Board on the other. 


H.—Correspondence relating to the adoption of the 
title “Royal Society of South Australia’”’ (1879-81). 
The minute books from September, 1853, to the end of 
1872, do not appear to have been preserved, but reports of 
the monthly meetings appear with a fair degree of regularity 
in the daily papers of that period. 
The first Annual Report was read on January 30, 1854. 
It consists of four pages (parliamentary size) and contains the 
personnel of the Council and list of members, together with a 
copy of Laws of the Society and a digest of its Transactions 
and Proceedings. Similar reports continued to appear until 
1858, when the month for the Annual Meeting was changed 
to July. No further printed reports were issued for some 
years after this, although they were evidently read and fully 
published in the newspapers. 


Annual meetings were again changed to August for 1859 
and 1860, and to October from 1861-3. Thereafter the year 
has apparently always closed at the end of September. 

Printed reports (in quarto form) re-appeared in 1865, but 
again ceased in 1872. 

A brief manuscript report for 1876-7 is to be found among 
the miscellaneous papers. From this date onwards they have 
been issued annually in their present form. 

It should also be mentioned, that brief abstracts of the 
_ Annual Reports of the Society, from 1863-84, are to be found 
_ as appendices to the Annual Reports of the S.A. Institute. 


a LE ee ee — 


620 


(b) Inception of the Society. 


The Society was founded on January 10, 1853. 

On the afternoon of that date five prominent citizens of 
Adelaide met at the house of Mr. J. L. Young, in Stephen’s 
Place, hardly a stone’s throw from this building, for the 
purpose of establishing a Society ‘for the discussion of all 
subjects connected with literature and arts.’’ 

John Howard Clark occupied the chair at this preliminary 
meeting, and there were also present: Messrs. J. L. Young, 
C. G. Feinaigle, —- Jones, and Dr. William Gosse. 

Three of these names, viz., that of J. H. Clark, a former 
editor of The Register; J. L. Young, the Principal of a well- 
known scholastic institution; and Dr. Gosse are still well 
remembered. 

Mr. Jones apparently did not attend any further meetings 
that year, and his name does not appear on the list of members 
published in 1854. His identity is probably lost in the mists 
of time. 

The fifth man, “)Charles Gregory Feinaigle, was the 
originator of the scheme, and as such claims our consideration. 
His residence in South Australia was of short duration, and 
consequently he is comparatively unknown in this State. I am 
indebted for much of the following information concerning 
him to the courtesy of the librarians of the Melbourne and 
Mitchell Libraries :— 

He was born in 1818, and graduated B.A., Trinity 
College, Dublin, in 1839. The date of his arrival in Adelaide 
is uncertain, but his name appears for the first time in the 
South Australian Almanac for 1851, as Headmaster of the 
High School, on the S.A Company’s premises, North Terrace. 
This was a proprietary school, apparently just founded, with 
shares at £5 each. J. L. Young arrived in October, 1850, and 
his first position was that of Assistant-master in the High 
School. In 1851 both these young men, were seized with the 
gold fever and went to the Victorian diggings. After an 
absence of several months Young returned, and was induced 
to open a school in Ebenezer Place, off Rundle Street East, 
but the movements of Feinaigle are not chronicled. From the 
facts already related, it is evident that he returned to 
Adelaide before the beginning of January, 1853. He occupied 
the chair, and read a paper on “The Mathematical Theory of 
Musical Harmony on April 25 of that year, and is mentioned 
in the First Annual Report, January, 1854, as ‘“‘being now ; 
absent from the colony.” For many years thereafter his 


_ (J. H. Clark in report of meeting of Philos. Society in The 
Register, 23/9/63. 


621 


name appears as a corresponding member, with a Melbourne 
address. As a matter of fact, he entered the Victorian Public 
Service as a clerk in January, 1854, and subsequently filled 
various positions in the Census, Police, and Mines Depart- 
ments. While still in Melbourne, he contributed a paper 
to the Society in September, 1863. He retired from the 
Victorian Service on a pension in 1877, and died at his resi- 


_ dence, South Yarra, after a long illness, on March 16, 1880. 


Three preliminary meetings were held at Stephens Place, 
and at these rules were drawn up for the government of the 
Society, and the annual subscription fixed at a guinea. 
Visitors were to be admitted to the meetings on introduction 
by a member, and they were allowed to take part in the dis- 
cussions, a privilege not infrequently exercised. It was 
decided that the election of members was to be by ballot, one 
negative vote excluding. It is worthy of note, that one can- 
didate was so excluded, during the first few months of the 
Society’s existence. 

In addition to such routine business, the roll of member- 
ship was greatly increased, and it is safe to say that the 
young Society already included within its ranks the best 
literary and scientific talent to be found in the city. Some of 
its members were men of undoubted ability and marked 
originality of character. A list of foundation members will 
be found in the Appendix, but it may not be out of place 
to refer more particularly to a few of them. 

Edward Davy, a versatile doctor, had a most extra- 
ordinary career. While still in England, he had already 
been recognized as a formidable rival to such men as Cooke 
and Wheatstone, in the new field of telegraphy. Not only 
was he an inventive genius in this science, but in many 
other branches as well. Quite suddenly, when his discoveries 
seemed likely to lead to wealth and eminence, he appeared to 
lose interest, and sailed for Australia as surgeon to an 
emigrant ship. Reaching the new colony in 1839, he aban- 
doned his profession and engaged in pastoral pursuits. Then 
followed a career of journalism, and for about three years he 
was editor of The Adelaide Examiner. Later still he became 
manager of the Yatala Copper Smelting Works. He retained 
this position for a few years, and then relinquished it in 
favour of the control of the Government Assay Office, where 
for the first time in Australia gold tokens were coined. Owing 
to his success in this department, he was lured to a similar 


- appointment in Melbourne at a salary of £1,500 per annum. 


Owing, however, to necessary retrenchments, his new appoint- 
ment was of short duration, and in eighteen months he was 


once more a farmer, this time coupled with the practice of 


622 


his profession as a _ sideshow. Finding that farming 
did not pay, he turned his attention to medicine and muni- 
cipal affairs in the sister colony of Victoria, and ultimately 
became mayor of a country town and a Justice of the Peace. 
He appears to have been an active member of the young 
Society, aud as a Corresponding Member retained his interest 
for many years after leaving Adelaide. 

Then there was Charles Mann, a former member of the 
South Australian Literary and Scientific Association, and at 
the period under review Crown Solicitor and a stalwart in- 
tellectual in the city. On account of his influence and 
literary tastes, he was an important accession to their ranks. 
He became their first Honorary Treasurer. 

J. L. Young never held office, but his intimate associa- 
tion with the infancy of the Society makes the picture incom- 
plete without him. For this reason, and also for the fact 
that we are partly indebted to him for that highly-finished 
product of his art, our immediate Past-President, I would 
like to see his portrait included in our family album. Owin 
to the good offices of a former pupil (Mr. F. W. Bullock), 
this portrait is forthcoming if the Council will accept it. — 
Unfortunately, owing to the lack of book-space, the walls of 
this room do not adapt themselves to the hanging of portrait, 
otherwise it would appear desirable to take immediate steps 
to secure as many photographs of foundation members for this 
purpose as possible. 

Young and Clark had been fellow-students at King’s 
College, London, where the former was educated as a profes- 
sional engineer. They were the same age, 23, when the pre- 
liminary meeting of the Society took place. 

John Howard Clark was undoubtedly the backbone of the 
Society and the most outstanding figure in its activities for 
upwards of twenty years. He was its first Hon. Secretary, a 
position which he held for nine years. Thereafter he became 
Hon. Treasurer, an office for which he was admirably adapted 
by reason of his early training as an accountant. His con- 
nection with The Register from 1865 onwards proved of 
the greatest service to the Society in the troublous years of 
financial depression, when the strictest economy had to be 
exercised in regard to printing. 

Another live wire among these pioneer members was W. W. 
R. Whitridge, journalist and pastoralist. He was editor of 
The Austral Examiner and subsequently of The Register. 
He early advocated the value of publicity, and the admission 
of -representatives of the Press to the monthly meetings— 
advice which the Society adopted with much profit to itself. 
He was a brilliant literary man, but unfortunately died at 
the early age of 36. 


} 


623 


The first ordinary meeting of the fully constituted Society 
was held on February 21, 1853, at the new City Council 
Chamber, then situated at the back of the present Town 
Hall. No rental was charged for the use of this room, and 
afternoon meetings were held there every month, until the 
close of 1858, when a change seems to have been made to 
White’s Commercial Rooms, where the Majestic Theatre now 
stands. Its work began to attract immediate attention, and 
resulted in the addition of such conspicuous men as the 
Governor of the State, the Chief Justice (Sir Richard Hanson), 
Sir George Strickland Kingston, Sir Arthur Freeling, Dr. 
Andrew Garran (another editor of The Register and subse- 
quently editor of the Sydney Morning Herald). The Press 
was well represented, and as a consequence the meetings 
received full and eulogistic reports in the daily newspapers. 

One of the earliest matters to receive consideration was 
the formation of a Museum to illustrate the natural history 
of the colony; but the difficulty which prevented the idea 
crystallizing into practical form, was the necessity of pro- 
curing suitable premises, and the high cost of rentals. It was 
suggested that the Government might possibly be willing to 
assign a room in one of the public departments for this pur- 
pose, and the matter was accordingly left in the hands of Mr. 
B. H. Babbage to privately sound the Minister before making 
formal application. Later on it was ascertained that the 
Mechanics’ Institute was contemplating a similar application, 
and although the matter was left in abeyance by the Society 
for the time being, it was never lost sight of, and frequently 
claimed their consideration at subsequent meetings. 

Towards the close of 1853 the rules which had been in use 
since the founding ofethe Society were reconsidered and a 
series of ‘‘laws’’ substituted. The chief alterations had to 


do with the formation of a Council or Executive body, 


Hitherto a Chairman had been elected to conduct the business 
of each meeting. Under the new laws, the officers consisted 
of a President, two Vice-Presidents, a Treasurer, and a 
Secretary. They were elected annually and together consti- 
tuted the Council. For a great many years after this, it 
became the custom to elect the: Governor of the Province to 
fill the chief position on the Council. There was only one 
exception to this rule, when, owing to the transfer of His 
Excellency Sir H. E. F. Young to Tasmania, B. H. Babbage 
was elected to the Presidency. 

At the end of the first year the membership stood at 
35. The young Society was now planted firmly on its feet, 
and in a position to state its objects and aspirations with a 


considerable degree of precision. They were twofold. (1) 


624 


“Tt was sought to afford an agreeable medium of intercom- 
munication to those whose tastes led them in pursuit of similar 
studies” ; and (2) to “‘present a means of illustrating and 
recording the many interesting phenomena, which are alto- 
gether peculiar to this country, and which it is feared will 
otherwise be lost in a very few years’ time, to the records of 
science.’’ In the papers contributed during that year, the 
second of these objects had not been overlooked, and at least 
two of them dealt exclusively with matters of local interest, 
that by Mr. M. Moorhouse on ‘‘The Structure of the Abor- 
iginal Dialects’? being of outstanding importance. 

The activities of the Society were not confined to the 
reading of papers, but extended to other matters affecting 
the public weal. 

With such APE ae) 23. eh. Babbage, Charles Bonney, 
and A. H. Freeling in their ranks, it is not surprising that 
a spirited interest should be aroused in exploration of the 
Province, important to them not only from an economic 
but also from a philosophic point of view. This was particu- 
larly the case in regard to the North-west Interior, which, 
like so many other parts of the continent, was then a terra 
mcognita. Rumours of various kinds had filtered through 


to them from native and other sources, which led them to — 


believe that this vast tract of country contained not only 
areas of pastoral importance, but also material of a scientific 
nature which intimately concerned them as a Society. In 
1854 a sum of £3,000 was passed by the Legislature for the 
purpose of exploring this portion of the colony, but owing 
to the difficulty of securing the services of a suitable leader, 
the sum still remained as an unexpended balance at the end 
of the financial year. . 

In 1855 the Society appointed a Special Committee to 
memoralize the Governor upon the urgency and importance 
of this enterprise, and also to collect such information as 
might tend to facilitate the organization and assist the oper- 
ations of an exploring party. 

In addressing His Excellency Sir R. G. MacDonnell, the 
memoralists pointed out, that ‘‘there exists at present a large 
and increasing demand for additional accommodation for the 
flocks and herds of the stockholders, occasioned partly by the 
rapid increase in the numbers of sheep and cattle themselves, 
and partly by the amount of land which has recently been 
taken up for agricultural purposes. That the very limited 
extent of the known and settled districts of the colony, coupled 
with the fact that the North-west Interior comprises an area 
of at least 150,000 square miles, offers reasonable ground for 
hoping that a knowledge of the character of this vast and 


i) oe 


——— NS 9 eee Beet ee er i 


625 


unexplored region would afford ample room for the extension 
of pastoral pursuits. That without pausing to dwell upon 
the desirability and importance of extending the limits of 
geographical science, the further consideration and knowledge 
of this extensive tract of country, may be reasonably expected 
to lead in an important degree to a greater development of 
our mineral resources, and thus open up new fields of enter- 
prise and additional sources of wealth to our colonists.’”’ To 
the memorial His Excellency gave a mest sympathetic reply 
and requested the Committee to submit to him a plan for 
such an expedition, together with an estimate of its probable 
cost; also to ascertain whether a suitable leader could be 
found to undertake the command of the party. 

A sheaf of correspondence between the Committee and 
such experienced bushmen as Dr. J. H.-Browne, J. McKinlay, 
E. B. Scott (a friend of Eyre), Price Maurice, and many 
others, shows that this request was promptly obeyed. As a 
result, we see the plan of the expedition outlined in a letter 
from the Committee to John Williams, of Black Rock, who 
_ had written, that if possible he would ‘‘be happy to undertake 
__ the business, notwithstanding the present discouraging aspect 
__ of affairs in that quarter.’”’ The plan was that an expedition, 
consisting of about eight men, should be landed at Fowler’s 
| Bay, make its way if possible due north to the north-east 
| corner of the Province, and return in a south-easterly line 

to the head of Spencer Gulf. In the letter they asked per- 

- mission from Mr. Wilhams to mention his name to His 
Excellency as a suitable leader. | 
Despite the energy displayed by the Committee, this par- 
ticular expedition did not eventuate, and the vote of £3,000 
was appropriated for other purposes. It is significant. how- 
ever, that the plan of Hack’s Expedition in 1857 was almost 
identical with that advocated by the Society two years pre- 
viously, except that it started from Streaky Bay. It is, 
therefore, probable that the Government was not altogether 
uninfluenced by the recommendation of the Committee. 

Hack’s effort was followed, in 1858, by another Govern- 
ment Expedition into the interior, under the leadership of a 
former President of the Society, B. Herschel Babbage. 

Another matter of a public nature, which claimed the 
_ early attention of the Society, was the establishment of a 
— South Australian Institute, which should be erected and 
. maintained by the Government, and should have for its 
- object the fostering of the arts, sciences, literature, and 
philosophy. It was thought that one of the functions of 
such an institution would be the establishment of a Natural 
History Museum, and that it would also provide accommodation 


a a 


626 


for such societies as might become incorporated with it. 
The Philosophical Society, during the first few years of its 
existence, was dependent on the goodwill of the City Council 
for a room in which to hold its meetings, and it eagerly 
desired something in the nature of a permanent home. 

The only existing institution having a somewhat similar 
scope was the almost moribund South Australian Library 
and Mechanics’ Institute. It was evident that this organiza- 
tion could not long survive by its own unaided efforts, and 
it was already, in 1853, seeking Government support to avoid’ 
extinction. It seemed probable that its purpose could be 
most satisfactorily achieved by its conversion into such an 
Institute as had been dreamed of by our Society. Conse- 
quently the two organizations joined issue in their attempts 
to secure this desirable object, but it was chiefly due to the 
energy displayed by the Society’s representatives, John 
Howard Clark and B. H. Babbage, that their efforts were 
ultimately crowned with success. 

In 1856 the South Australian Institutes Act was passed, 
and within a month the Institute began its career of useful- 
ness. It was administered by a Board of six, three of whom 
were appointed by the Governor and three by the Societies 
which it had power to incorporate. : 

The Act provided that a sum of not less than £500 should 
be made available for maintenance, and a short amending 
Act enabled the Board to make advances of money to Incor- 
porated Societies. No time was lost by the Society in making 
its application for incorporation, but although a sympathetic 
reply was received, it was nevertheless pointed out, that 
however desirous the Board of Governors might be to effect 
such incorporation, the circumstances in which they were 
temporarily placed rendered an immediate junction imprac- 
ticable. The reply had reference to certain defects in the 
Act which required amendment, and also to the difficulty in 
regard to accommodation. 

The Institute was at this time housed in Green’s 
Exchange Buildings, but an endeavour was being made to 
secure more commodious premises. Owing to the difficulty of 
obtaining suitable rooms, the Board at length decided to apply 
to the Government for the erection of a building on public 
land. It was estimated that the cost would be approximately 
£4,000, and immense energy was displayed by all parties to 
induce Parliament to make this sum available for the purpose. 

Not the least active participant in these proceedings was 
the Philosophical Society. Among the papers preserved in 
the Archives is a draft memorial to the House of Assembly, 
in the handwriting of John Howard Clark, who was then 
Hon. Secretary. 


a 


WI 
\ 
1 


———— 


SARE TI. Pew ae 
See Pow 


627 


Inter alia this memorial states :—‘‘Your memorialists are 
authorized to negotiate for the incorporation of the Adelaide 
Philosophical Society with the South Australian Institute. 

. It is impossible that such incorporation can be 
effected, until the S.A. Institute has at its disposal a per- 
manent and suitably designed building; affording ample 
accommodation for its various requirements. One of the 
principal objects of our Society is the formation of a Museum 
illustrative of the Natural History of the Province, and it is 
useless to take any steps in furtherance of this object until 
a suitable room is provided for preserving the specimens 
collected, although valuable specimens would then be imme- 
diately available, many of which in a few years’ time it would 
be impossible to replace.’’ 

Parliament found it impossible to resist the pressure 
brought to bear upon it and the sum was passed. 

The next matter that aroused great controversy was the 
site of the proposed building. Parliament had selected a site 
between the back of the present City Baths and the Cheer-up 
Hut. This evoked the most heated discussions in the news- 
papers, and resulted in many deputations and public meetings. 

The Philosophical Society threw the weight of its in- 
fluence into the scales on behalf of a more prominent and 
accessible position. In addition to much private wire-pulling, 
they embodied their views in another memorial to His Excel- 
lency Sir R. G. MacDonnell, at that time their President. 
Once more the well-known caligraphy of J. H. Clark can be 
recognized in the draft. “‘Your memoralists have learned 
with regret that it is proposed to erect a building, to be 
devoted to the objects of the Institute, in a locality which 
appears to them to be objectionable in many respects; inas- 
much as the site selected is so much lower than North Terrace, 
that not only will the building (which should ultimately become 
one of the chief ornaments of the city) be almost hidden from 
sight, but its situation will be neither convenient for public 
access, nor advantageous for meteorological observations, 
whilst the steep gradient of the City Bridge Road will neces- 
sarily render the approaches to the building unsafe for the 
large number of vehicles, which will hereafter be frequently 
gathered together at night, on the occasion of lectures or 
soirees connected with the Institute or its affiliated societies. 

Inasmuch as the Houses of Parliament are, and 
long will be, amply sufficient for the requirements of the 
colony, it is “needless to leave unoccupied the excellent site 
for an important public building, which could be made avail- 


able at the corner of North Terrace and the City Bridge Road, 


and which is at present said to be preserved for future new 


628 


Houses of Parliament. Your memoralists pray that Your 
Excellency will be pleased to direct, that the building for 
the S.A. Institute may be erected upon the site last men- 
tioned, or upon some other site better suited to the present 
requirements and ultimate importance of the Institute than 
that now in contemplation.” 

Once again the faith of the Society in memorials was 
justified. Parliament meekly bowed its head before the storm 
of public protest, and the proposed site was changed to that 
occupied by the Institute to-day. 

In this, as in all public matters which touched its objects 
or its principles, the Society was ever ready to fight for the 
common interests of itself and its friends. Impecunious as it 
was, 1t possessed in great measure the brains of the small 
community, an endowment of greater importance and influence 
than mere material wealth. It was instinctively alert to 
recognize the value of powerful friendships, such as that of 
the Governor of the colony, and, above all, it fully realized 
and appreciated the power of the Press. 

Under the Act, the subscribers to the Institute exercised 
the privileges of an incorporated Society, and at the first 
annual meeting in October, 1857, they rewarded the services 
of J. H. Clark by electing him as their representa- 


tive on the Board of Governors, an undoubted honour for so 


young a man. 

The Society’s tenacity of purpose was one of its most 
valuable assets. In its early infancy it had desired a Museum, 
a desire constantly foiled by almost insuperable difficulties 
and only realized when passing into middle age. But until 
realized its purpose was always in evidence. Persistent 
flagellation of the public interest, as well as that of the 
Governing Board, kept the matter alive, or at least in a state 
of suspended animation. 

In its first report, the-Board speaks hopefully of the early 
accomplishment of this object. ‘‘As regards a Museum, the 
prospects of the Institute are most satisfactory. Extensive 
collections of great and varied interest await only a room 
for their reception. The proprietors of mines in this colony 
have in all instances complied with the request of the 
Governors to be furnished with specimens characteristic of 
their various properties, so that at its opening the Museum 
will exhibit an epitome of the mineral riches of South Aus- 
tralia. To the Directors of the Burra Mine, the Governors — 
are indebted for a very extensive and interesting collection 
just lately come to hand. Many valuable presents are also — 
promised by private individuals. His Excellency Sir George © 
Gray writes from Cape Town, in reply to an application made 


629 


on behalf of the South Australian Institute, that he has 
directed many interesting specimens to be collected and for- 
warded hither, including a complete series of the copper ores 
of the colony over which His Excellency presides.’’ 

‘‘The Governors believe, that when the Museum is estab- 
lished many other additions to its contents will be received 
from abroad, and that many of our own colonists, who are 
known to possess miniature museums, will be anxious to 
incorporate them in the public collection.’’ 

In their second report, October, 1858, they acknowledge 
receipt of the collection presented by Sir George Gray and 
also the purchase of ‘‘a very interesting and extensive col- 
lection of shells.”’ 

It will be seen from the tenor of these reports, that the 
institution at this time in the minds of the Governors was 
almost entirely a mineralogical collection, and very far 
removed from the Natural History Museum, which was an 
early objective of the Society. Nevertheless, it indicated a 
slight advance in the right direction. 


(c} Incorporation with the S.A. Institute. 
Es Incorporation -of the Philosophical Society with the S.A. 
| Institute was duly effected in October, 1859. The terms, 
| however, were less favourable than the Society had antici-. 
| pated. It was to receive certain clerical services from the 
Institute, but a room for its exclusive use was refused and 
| accommodation was guaranteed for monthly meetings only. 
For the privileges of incorporation, the Society was to con- 
tribute one-third of its gross annual income, but the minimum 
contribution was fixed at £15. In a letter to the Society, 
of which he was still Secretary, Mr. Clark carefully points 
out that “‘the sum required is not so much im the nature of 
an amount paid away for house rent and clerical services, as. 
a contribution towards a fund to be expended for the general 
benefit of all connected with the institution, and in the ex- 
penditure of which the Society will have a voice.’’ He illus- 
trated his meaning by the statement that the Governors had 
already sent to England for a valuable microscope and a pair 
of 36-in. globes, and that doubtless with accession of funds 
the stock of philosophical apparatus would be speedily 
_ increased. 
. It is to be feared that in the years that followed the Society 
too often lost sight of this statesmanlike view, when the hand 
of adversity pressed heavily upon it. : 
‘ The Society was, of course, now entitled to elect a repre- 
_ sentative Governor, and its first choice fell upon B. H. Bab- 
_ bage, who was a foundation member and its President from 


630 


1855-6. Another of its most active members, Mr. Whitridge 
was also elected to the Board by the Society of Arts, so that 
with three of its prominent members on the Executive of 
the Institute, its interests would appear to be well protected. 


A delay arose in the erection of the building, and mean- 
while the Institute continued to occupy the premises in 
Green’s Exchange and the Philosophical Society those in 
White’s Commercial Rooms, King William Street. 

The new building was opened with great ceremony by 
His Honor Sir Charles Cooper on January 29, 1861. The 
room allotted to the Society was upstairs, immediately over 
the lhbrary, with a south-easterly aspect. Its floor has re- 
cently been removed to increase the shelving accommodation 
for books. ae 

The Museum consisted chiefly of a mineralogical col- 
lection, and occupied a long narrow room running across the 
upper story at the rear of the building. There it remained 
under the curatorship of F. G. Waterhouse, naturalist in 
McDouall Stuart’s Expedition, for a period of twenty years. 
During all these years, very little expansion was possible, 
owing to the lack of space, and it was not until it was removed 
to more commodious premises that any serious attempt to 
form a zoological collection could be entertained. 

Incorporation with the 8.A. Institute was certainly not 
followed by the signal advantages hoped for by the Philo- 
sophical Society. In many ways it proved a grievous dis- 
appointment. 

In the first place, the limited accommodation afforded to 
the long-desired Museum had to a great extent shorn that 
institution of its utility. In no sense could it be regarded 
as a collection representative of the natural history of the 
colony, and it was therefore of little value as an attractive 
and popular set-off to the technical aspect of the Society’s 
work. In this matter the Board was powerless to help, not 
that it was lacking in sympathy, but merely because the build- 
ing was altogether too small for the purposes_to which it had 
been dedicated . 

Then, again, the Society had hoped to derive some finan- 
cial advantage from the union. The Amending Act gave 
the Board power to advance moneys to incorporated societies ; 
and the Consolidated Act of 1863 also gave it discretionary 
power to make a grant to any Scciety so imcorporated. 

In addition to the annual subsidy made by the Govern- 
«ment to the central institute, sums of varying amount were 
also paid to it for allocation among affiliated country institutes. 
It was clearly the intention of the Legislature, that the 
Philosophical Society as an incorporated body should benefit 


: 631 
under the Parliamentary vote, yet the Estimates were so 
worded that it was precluded from this privilege. 

It was not until many years later (1878) that this dis- 
ability was removed, and it was placed on an equality with 
country institutes, receiving, like the latter, an annual 
Government subsidy equal to the subscriptions for the current 

ear. 
j Thus for nearly twenty years it paid to the Institute 
out of its small income of about £50, a sum which it regarded 
as practically a rental, for the empty privilege of electing 
a Governor, who was powerless to promote its interests or 
adjust its grievances. 
It is suggestive of its poverty, that for several years after 
__ incorporation the publication of the brief annual reports sud- 
_ denly ceased. The balance-sheets show, that since its founda- 
tion the Society had been striving to establish a reserve fund, 
| which in 1858 stood at £77. This sum would have more than 
. sufficed for the continuation of the reports, but it is probable 
that in view of an uncertain future, it had been decided to 
temporarily discontinue them and depend for publicity upon 
the goodwill of the newspapers. Such strict economy may also 
_ have been prompted by a desire to assist in the furnishings 
, of the Museum, about which much difficulty had unexpectedly 
arisen; for it is on record that in 1861, the substantial sum 
of £50 was expended for this purpose. 
' At this period of its history, the Society was extremely 
isolated from the scientific world, having no publications to 
exchange for the Proceedings of other bodies and having 
practically no funds wherewith to effect purchases. As early 
as 1863, the position had indeed become so acute that it clearly 
_ contemplated secession, and was only deterred by the fear of 
losing its property. In a letter to the Board dated June 15 
of that year, the Council wrote requesting it to confer with 
their Special Committee as to the interpretation which should 
be placed upon certain clauses in the terms of incorporation. 
It may be here pointed out, that the Board had retained the 
right to place its own construction upon any debatable 
terms in this agreement. The letter proceeds: —‘‘The Com- 
mittee particularly desires to know the views of the Board, 
under the possible circumstances of entire removal of the 
Philosophical Society to other premises. Also upon the 
restraints which the terms of incorporation have upon the 
Society ; its rights to remove, sell, exchange, or otherwise dis- 
_ pose of its movable property, such as specimens and instru- 
ments. Also upon any independent power which the 
_ Governors may claim to exhibit, remove, lend, or use any of 
_ such property without previously obtaining the consent of the 


632 


Society. -Also seeing, that by the Act, the Governors have 
power to make bye-laws, whether such bye-laws would alter 
the interpretation now given, relative to the terms of 
incorporation.”’ | 

The letter was not wholly without guile. If they could 
remove their property to other premises, without fear of con- 
fiscation, then the path would be open for any future course 
of action they might desire to pursue. 


The reply from the Board, however, was guarded, and 
in the nature of a compromise which did not materially 
improve their position. It announced, that as a result of the 
conference, ‘‘the Board are willing to modify the articles of 
incorporation between the S.A. Institute and the Philosophical 
Society, so as to make them accord with the incorporation 
clauses of the schedule of Statutes and Rules.’’ This meant 
that property could only be removed with the consent of the 
Governors, and with this proviso, should vest in the S.A. 
Institute, only in the event of dissolution of the incorporated 
Society. 

The schedule of the Acts of 1855-6 had provided that 
property accumulated by incorporated societies should become 
vested in the Institute forthwith and without reservation. But 
this was a schedule which the Board had power to alter or 
amend, subject to the approval of the Legislature, and which 
had already been so amended in 1861. 

A new Consolidated Act, embodying the undertaking of 
the Board, duly received assent the following November. 


The room in which the Society held its meetings was | 


only at its disposal for twelve meetings a year, whereas the 
actual number of meetings exceeded this. It was inade- 
quately furnished, and it was therefore necessary to incur 
the expense of erecting a cupboard and shelving, of which the 
Society had to bear half the cost. 

The one bright spot in the landscape at this period was 
due to the unvarying kindness and sympathy received from 
the Press. Although the Society had no means of communi- 
cating its work to the scientific world, its members continued 


to read their papers, and the Press, out of the greatness of 


its charity, continued to publish them. 


After reading the reports of the 8.A. Institute, one is 5 


forced to conclude that the difficulties which confronted the 


Philosophical Society after its incorporation were not the 


callous creation of the governing body of that institution. 
The very constitution of the Board precluded a charge of in- 


difference. Even the Secretary (Robert Kay) was a founda-— 


tion member of the Society and a man of strong personality. 


Pts ae vs 


633 


The fact was, there was no period in the history of the 
institution when it had money to burn. The Act was per- 
missive, and when the loudly-voiced demands of the subscribers 
and country institutes had been met, there were no funds left 
in which the Society might participate. 


Nevertheless, the Society continued to do good work; 
many of its papers were of high quality, and it continued 
to interest itself in economic matters of public utility. 


In 1862 it attempted to establish an Acclimatization 
Society. For this purpose it appointed a Committee to carry 
out this .object. Much information was collected and an 
important paper was prepared and read by Mr. G. W. Francis, 
an enthusiastic advocate of the movement. This paper was 
published by the Society, and duly circulated, but did not 
create sufficient public interest to warrant further steps in 
regard to the matter. About a year later, however, such a 
body was duly founded in Adelaide, the success on this occa- 
sion being undoubtedly due to the earlier efforts of the 
Philosophical Society. 

Another matter which claimed its serious attention in 
1866 was that of the city drainage, a subject of great import- 
ance to Adelaide from the standpoint of the health and com- 
fort of its citizens. A series of resolutions were formulated 
and embodied in a memorial to the City Council, with the 
result that a Bill was introduced into the Legislature to 
enable the Corporation to initiate a modern system of sani- 
tation. 

In the same year, the question of railway gauge exercised 
the minds of its members and formed the subject of much 
interesting discussion. The result of this was the adoption of 
several resolutions, one of which was: ‘“That the saving in 
the construction of a 3 ft. 6 in. line over that of a 5 ft. 3 in., 
calculated for a similar amount of traffic, would be by no 
means proportionate to the difference of width of gauge, and 
that our branch lines should be constructed as lightly and 
as cheaply as possible with a 5 ft. 3 in. gauge and worked 
with horse-power, until the amount of traffic renders the use 
of a light locomotive at low speed more economical.” 


Towards the close of 1870, there had been held nearly 
170 meetings at which upwards of 200 papers had been read. 
Many of these dealt with geographical exploration and 
branches of applied science relating to horticulture, metal- 
lurgy, and meteorology; others were important contributions 
to the geology and natural history of the Province. 


John Howard Clark became Hon. Treasurer in 1863, a 


' position which was not without its penalties. In 1866, the 


- 


634 


financial statement shows a small balance of £1 16s. 10d. due 
to the Treasurer. The annual exhortations by the Hon. Secre- 
tary requesting the members to promptly meet their 
obligations do not appear to have met with a ready response, 
for the following year the subscriptions had still further fallen 
off, and the amount due to the Treasurer had increased to £30. 

The Council recognized two possible ways of adjusting 
this unsatisfactory state of its finances. The first was by 
curtailment of expenses, which could not be ‘done without 
diminishing the usefulness or impairing the attractiveness of 
the Society ; the other way was by a material increase in its 
membership. The second alternative was chosen as a solution 
of the difficulty, but it evidently failed, for in 1868 it 
was found necessary to realize upon the property. This 
was done by effecting a sale of some museum cases to the 
S.A. Institute for a sum of £40. As these same cases had 
cost them £50 a few years previously, the transaction could 
not be regarded as altogether a favourable one. It enabled 
the Society, however, to restore the Treasurer’s account to an 
equilibrium, leaving it with a small credit balance for 
the year. This balance was increased the following year, 
by a still further encroachment upon the property. This 
time it was a sale to the Institute of the valuable set of 
Transactions of the Royal Society of London, for a sum 
slightly in excess of the original cost. The proceeds of the 
sale were received in two annual instalments. The cost of 
printing the yearly report and transactions for the succeeding 


year was, however, greatly in excess of what had been pre-— 


viously paid, so that 1870 ended with a much more slender 
balance than had been anticipated. 

The colony had now failen upon troublous times, and 
these were reflected in the fortunes and finances of the Society. 
In its report for the two years ended September 30, 1872, 
the Council, while expressing its satisfaction at the growing 
interest in its proceedings by the public, as exhibited in their 
general attendance at its meetings, nevertheless directs atten- 
tion to the fact, so significant of the times through which it 
was passing, that out of 62 ordinary members, only 34 had 
paid their subscriptions for the current year. 

It is not therefore surprising, that a letter dated 8/2/71 
should be found amongst the correspondence in the Archives, 
asking the §.A. Institute to reduce the contribution to £12 
per annum. Two reasons are advanced in support of this 
request: (1) ‘“‘Because we can secure a comfortable room in 
the Town Hall for 10s. 6d. per meeting,’’ and (2) “‘because 
our expenditure is in excess of our income.” The request 
was granted. 


635 


The time had evidently arrived for drastic retrenchment, 
and no further reports of transactions were published for 
five years. 


(d) Establishment of the University and the 
Coming of the Grant. 


Although the prevailing depression in the colony in the 
early seventies had robbed the Society of that vigour which 
had characterized it in previous years, two all-important 
events were about to arouse it into unprecedented activity and 
usefulness. These were the establishment of the ey 
and the coming of the Government grant. 


The influence exerted by the University can hardly be 
over-estimated. It did not intrude itself gradually, but was 
almost cataclysmic in its suddenness. The Society began to 
radiate vitality. The scientific leader for which it had waited 
so long appeared in the person of Professor Ralph Tate, whose 
energy was tremendous. Popular lectures were delivered, 
public interest was aroused, resulting in a sudden accession 
of strength to its ranks and a welcome increase in its funds. 
Effort and knowledge now became organized, and the Society 


| was soon raised from a mere parochial body to an assured 


place in the scientific world. . 


The following year the publication of its Transactions 
was resumed and appeared in a form more in accordance with 
those issued elsewhere. 


Throughout this new phase of development there was, 
however, always present the lurking fear of adversity. At all 
costs some permanent source of income must be discovered, 
to assure them against any further breaches in the continuity 
of their publications. In 1879 the Council once more 
approached the Board with a view of securing some financial 
benefit under the Institutes Act. Acting on the advice of 
the Board, and with the promise of its assistance, an applica- 
tion was made to the Government, asking that the Society 
should receive the same measure of support as had hitherto 
been accorded to the country institutes. The application 
was favourably considered, and resulted in an annual subsidy 
on their subscriptions. The subsidy for the year in question 
amounted to £118, which, though it did not enrich them, 
at least defrayed the cost of their Transactions and left a few 
pounds over for emergencies. 


Since then the Society has never looked back. Its publica- 


"tions have been continuous for forty-five years, and are now 
_ to be found on the shelves of every important scientific library 
_ throughout the world. 


636 


(4) Tue Roya Socirry oF SourH AUSTRALIA. 


(a) How we became “‘ Royal.’’ 


At the close of the seventies it became apparent to the 
Society that its investigations would receive added weight 
and dignity if it could include in the title a warrant of Royal 
favour. In other words, it appeared that there was something 
in a name. 


The occasion was an unusual one, and it was not known 
to the Council what procedure should be adopted to solicit the 
patronage of its Sovereign. There were at that time in 
Australia three Societies that rejoiced in the title of ‘“Royal’’ ; 
one in New South Wales, one in Victoria, and in Tasmania 
one that Tae ee founded many years previously by Sir John 
Franklin. Walter Rutt, who was then, as now, Secre- 
tary of the nee accordingly wrote to the ‘Secretary of the 
latter body, quiring as to the steps which had been taken 
when they had solicited a similar privilege. Dr. Agnew 
replied, that it had been necessary to make a search through 
the records in the Colonial Secretary’s Office, and subsequently 
through His Excellency’s despatches, for the purpose of 
obtaining specific information on this subject. Even then his 
search had been unsuccessful, but he had learned from Mr. 
Hull, a corresponding member of the Philosophical Society, 
and in 1843 a confidential member on the staff of Sir Eardley 
Wilmot, that on the occasion in question a despatch was sent 
to the Secretary of State, conveying a request that Her 
Majesty would be graciously pleased to become a Patron of 
the Tasmanian Society. At the same time, the claims of the 
Society to this mark of favour were duly set forth. A favour- 
able reply had eventully been received from the Home Govern- 
ment, and it had since enjoyed the title of “‘Royal.’’ A 
subsequent letter from Dr. Agnew conveyed the information 
that the despatch from Lord Stanley, Secretary for the 
Colonies, had been found. It stated that Her Majesty had 
graciously consented to become the Patroness of the Tas- 
manian Society, and had acceded to the request that it should 
be permitted to use the title ‘‘Royal.’’ 


Acting on this precedent, the Council of the Adelaide 


Philosophical Society addressed the following request to Sir 


Wilham Jervois, then Governor of the colony :— 

‘‘The Adelaide Philosophical Society, knowing that there 
is a wide and comparatively unexplored field for scientific 
research in this extensive Province of South Australia, is 
appealing to all who take an interest in such matters, to 
forward to the Society the results of their observations and 
investigations, in order that they may be collated and placed 


ee 


| 
| 
i 
| 
. 
; 


637 


on record for the benefit of the scientific world. The Council 
feels that a more ready response would be made to this appeal, 
and that more attention would be given by men of science 
throughout the world, if Her Majesty would graciously extend 
to it her patronage. In this view members of the Society 
concur. 


“T am therefore instructed to request you to kindly 
take the necessary steps to lay before the Queen the prayer 
of this Society, that Her Majesty will graciously consent to 
become the Patron of the Society, under the title of ‘The 
Royal Society of South Australia,’ and thus place it upon an 
equality with the Royal Societies existing for similar purposes 
in other Australian colonies. 

“Tf Her Majesty should be pleased to accede to this 
request, you would perhaps, as representative of the Crown 
in this Province, not object to accept the position of Vice- 
Patron. 

“The Society has done a considerable amount of work, 
and is desirous of widely extending its operations in the 
future.”’ 


His Excellency, a punctilious observer of the formali- 
ties, in his reply submitted the following suggestions for con- 
sideration by the Society :— 


1. That the Society’s application might conveniently take 
the form of a memorial to Her Majesty, and if on parchment, 
should be accompanied by a copy on folio paper. It should 
bear the signatures of the President and principal officers of 
the Society. 


2. As to matter, it would probably be of advantage, if 
the memorial contained, after the opening statement, a con- 
cise sketch of the origin and progress of the Society, of its 
funds, numbers, times of meetings, circle of subjects hitherto 
embraced, and transactions generally. Something should be 
stated as to the results attained. 'The memorial should also 
be accompanied by four copies of all printed matter relating 
to the proceedings of and subjects treated by the Society. 

He pointed out that the application was in a measure 


on the same footing as that recently made by the University 
of Adelaide for a grant of Letters Patent, and, as in that 


| case, a clear statement of the nature and position of the 


Society is requisite to obtain the object in view. 
' A memorial embodying the above suggestions having been 


_ submitted to His Excellency, he approved of its form, but 


_ deemed it advisable, as a preliminary step, that it should be 


i 


adopted at a full meeting of the Society. 


638 


The following memorial was at length forwarded by His 
Excellency :— 


‘To the Queen’s Most Excellent Majesty. 
‘‘May it please Your Majesty. 

‘“‘We the undersigned, acting on behalf of and at the 
request of the Council and members of the Adelaide Philo- 
sophical Society, as expressed by a resolution passed at a 
meeting of the Society on the 4th day of May, 1880, humbly — 
lay before you our prayer, that Your Majesty will be graciously ° 
pleased to become the Patron of the Society under the title 
of ‘The Royal Society of South Australia.’ 


“The Adelaide Philosophical Society was founded in 
1853, for the diffusion and advancement of the Arts and 
Sciences, by the meeting together of the members, for the 
reading and discussion of papers connected with the above 
subjects and by other approved means; and was, in 1863, 
incorporated with the South Australian Institute, under the 
provisions of the South Australian Institute Act of that 
year,@) which retained to such incorporated Societies their 
individuality and full independence of action. | 


“The Society has done much in the past to keep 
alive in a struggling and young community the im- 
portance of scientific research. The rapid extension of 
agriculture in districts which were until recently occupied 
only for pastoral purposes, the successful journeys of many 
exploring parties, the consequent advance of pastoral settle- 
ment in the interior and in the Northern Territory, the devel- 
opment of the country by railways and telegraphs, and con- 
sequent prosperity of the colony, have given increased 
opportunities and leisure for the collection of facts in the 
natural history of the Province, which has hitherto been a 
field almost unexplored by the scientific observer. The Society 
has, therefore, established correspondents throughout the 
Province, and the value of the results thus obtained and for- 
warded to the leading scientific societies of the world will 
be seen by an inspection of the two volumes of the New Series 
of the Society’s Transactions, copies of which are forwarded 
for Your Majesty’s information. The Old Series of Trans- 
actions, published only for distribution amongst the members, 
is now out of print. 

‘‘The Society, numbering at present about 110 members, 
meets monthly, and its income this year will exceed £200. 


(2) The year of incorporation was 1859. 


(3) The Institute Act was passed in 1856, the Amending Act 
the same year. 


639 


“It will be seen that the objects sought, and the results 
obtained, by the Adelaide Philosophical Society are similar 
and equal to those sought and obtained by the Royal Societies 
of New South Wales, Victoria, and Tasmania, and we feel 
that the Society will be largely assisted in its efforts to increase 
the value of these results, and that more attention will be 
paid by scientific men throughout the world, to the facts 
_ recorded year by year in its Transactions, if Your Majesty 
will be graciously pleased to accede to the request of your 
memorialists, who will, as in duty bound, ever pray, etc. 
‘“‘Ravtpx Tare, President. 
‘“‘WREDK. CHAPPLE, 
“‘Cuas. Topp, 
‘“‘THomas D. Smeaton, Hon. Treasurer. 
““WaLTER Rutt, Hon. Secretary.” 

Four months later a letter was received by the Governor, 
enclosing a copy of a despatch received from the Secretary ot 
State for the Colonies :— 

“South Australia. Downing Street, 
“No. 24. 3rd August, 1880. 

‘“‘Sir—I duly received your despatch No. 31 of the 15th 
May last, and submitted to Her Majesty the Queen the 
memorial from the Adelaide Philosophical Society, praying 
that Her Majesty might be pleased to become the Patron of 
the Society, under the title of the Royal Society of South 
Australia. | 

“T have now the honour to request that you will inform 
_ the President of the Society, that Her Majesty has signified 
her gracious approval of the Society being styled the Royal 
Society of South Australia. 

‘“‘T have the honour to be, etc., etc., 
‘“KIMBERLEY.”’ 


\ Viee-Presidents. 


| The final letter in this correspondence is from the Private 
Secretary to the President of the Royal Society of South 
Australia, dated 20/1/81, informing him that His Excellency 
wishes that the future volumes of the Royal Society should 
be forwarded to the Secretary of State for the Colonies, 
_ through him, and requesting the President to transmit to 
_ him three copies of the volumes in question, as published. 


(b) New Buildings and a New Act. 
: Almost from the beginning it was recognized that the 
accommodation in the S.A. Institute was inadequate for the 
_ purposes of a Museum. This became more and more apparent 


i 


640 


as time went on, and the restrictions which the limitations of 
space placed upon the collection rendered it practically value- 
less for natural history purposes. Not only was accommoda- 
tion insufficient for Museum purposes, but pressing need for 
expansion was also felt by the library and other departments 
under control of the Board. Similar inconvenience was ex- 
perienced by the incorporated Societies, and the Philosophical 
Society, whose meetings were now open to the public, found 
their room uncomfortably crowded, when any subject of 
special interest was being discussed. 

It became almost painfully evident at the end of the 
sixties that a new building was an urgent necessity, and in 
1871 Parliament, without dissentient voices, expressed its 
sympathy with the proposed enlargement. The Board recom- 
mended the erection of a new building to the east of the 
Institute, to consist of two wings, of which the western one 
was to be proceeded with first. 

Money was found by the Government for this purpose 
and the foundations were duly laid down in 1873. There 
seems, however, either to have been a lack of unanimity with 
regard to the proposed scheme, or else a desire on the part of 
the Government to delay public expenditure, for at this stage 
building operations ceased and a Royal Commission was 
appointed to inquire into the whole matter. 

The Commission favoured the idea of a Public Library 
and Museum to replace the Institute, and recommended that 
these should form two wings of a new building, to be erected 
to the east of the latter. No attempt was made to carry 
out this recommendation until 1876, when it was discovered 
that the foundations which had been laid three years pre- 
viously were unsound. New ones were laid, but these were 
again temporarily abandoned for two years, when it was found 
that they, too, had to be taken up and replaced. The founda- 
tion-stone of the present Public Library was ip lr, laid 
dain Gei/f.3 

This time the erection of the new building proceeded 
rapidly and without further delay. The western wing (now 
the Public Library) was sufficiently advanced for occupation 
in 1882, and as the Institute rooms occupied by the Museum 
were required for the new School of Design, the collection” 
was withdrawn from public view, and placed in the crypt and 
two smaller rooms of the new wing in the early part of that 

ear. 
Here it was submitted to a critical examination by Dr. 
Haacke, the new Director. This gentleman on his arrival 
stated his views very frankly as to what he considered the 
S.A. Museum ought to be. ‘‘In the first instance,’’ he says, 


Foe = 


641 


“there should be in South Australia no institution rivalling 

the Museum under my care, as this would not tend to further 

the scientific and educational interests of the colony. In Ade- 

_ laide we enjoy the existence of a Botanic Garden, with a 

_ Museum of Economic Botany, and of a University with a 

- museum for lecture purposes; a Zoological Garden is now to 
be established, and we shall have a Technological Museum in 
course of time. There are also small museums connected with 
some of the country institutes, and the Royal Society is 
endeavouring to promote the intellectual and_ scientific 
advancement of the colony. In the interest, not only of the 
Museum, but of all the above institutions, I respectfully beg 
you to take the following suggestions in the spirit in which 
they are made. 

“T think it would be wise to exclude any technological 
and botanical collections from the present Museum, where, 
however, all objects of zoology, ethnology, mineralogy, and 
geology should be gathered, as long as it is not thought advis- 
able to have special museums for each of these branches of 
science. 

‘“‘In the Zoological Gardens to be established, only living 
animals should be kept, and the museum in connection with 

_ the University should only contain such collections as will 
| be useful in lectures. Again, the country museums should be 
_ satisfied in having only good educational collections, while 
| all objects of scientific value should go to the central institu- 
tion, which, in connection with the Botanic Gardens, the 
| University, and the Royal Society, ought to represent science 
in South Australia.”’ 
| He then proceeds to outline the manner in which, in his 
4 


_ opinion, the collection should be displayed. . 

i} I think there are very few of these suggestions which 
would fail to meet with approval to-day. 
1} Two years elapsed before the collection was considered 
_ to be sufficiently advanced for public exhibition. A portion 
_ of it was then displayed in the northern half of the present 
| Library, which remained its home for ten or eleven years. 

| In 1894, that portion of the building occupied by the 
Museum was urgently required for Library purposes, and 
| the collection was once more removed, this time to the present 
' brick building which connects the two wings of the institution. 
Ri It is within the recollection of most of you, that at quite 
a recent date further room for expansion was urgently needed, 
and most of the eastern wing was appropriated for that pur- 
a. Then only were the early dreams of the Philosophical 
‘Society almost realized, by the existence in this city of a 
Museum containing an excellent and representative collection 


642 


of the natural history resoures of the State, and among other 
things the finest Australian anthropological and ethnological 
collection in the world. 

Even now the buildings are overcrowded, and there are 


many important specimens which have to be stored until 


space can be found for their exhibition. There are also gaps 
in the collection, some of which at this late period in our 
history we cannot hope to fill, but there are likewise desiderata 
which we still hope to receive from the hands of some diligent 
and patriotic collector. 

At all times the most cordial relations have existed 
between our Society and the scientific staff of the Museum. 
They have, I believe, in every instance been amongst the most 
honoured and respected members on our roll. Since 1884, 
when the Museum first became an institution worthy of the 
name, a representative of the Council has always been Chair- 
man of its Committee, a position which Professor Howchin 
has honourably discharged for the past twenty years. One 
can have no hesitation in saying, that it is now fulfilling the 
high scientific functions for which it was established and 
towards this success it would not be immodest for the Royal 
Society to claim some degree of credit and responsibility. 

The passing of the Public Library Act of 1884 created 
a somewhat curious situation, which at first appeared to 
threaten the interests and stability of the Royal Society. 

This Act, which abolished the S.A. Institute and super- 
seded it by a Public Library, Museum, and Art Gallery, 
renewed the privilege of the Society to elect a representative 
Governor cn the Board, but unfortunately it did not make 
provision for the incorporation or affiliation of societies. 

Without such provision the alarming fact became dis- 
closed, that the Government grant which the Society had now 
enjoyed for several years had quite suddenly lapsed, and the 
Society had therefore incurred liabilities during its financial 
year which there might be no means of meeting. 

It was a very anxious and perturbed Council that opened 
up a correspondence with the Board on this all-vital question 
in August of 1884. In reply to their inquiry as to how the 
position of the Society and the Government subsidy would be 


affected by the new Act, they were informed that the Act had. 


put an end to the S.A. Institute in the previous June, and, 
of course, at the same time to the incorporation with it of 
the Royal Society. Further, that the Estimates for the 
current year provided grants to country and suburban insti- 
tutes and also to affiliated societies; that as it might not be 
advisable to alter the wording of the line on the Estimates, 
the safest course for the Board to pursue would be for it to 


¢ 


| 


a ee 


7 


| 


643 


pass a formal minute declaring the Royal Society to be a 
Society affiliated to the Public Library, Museum, and Art 
Gallery. This, however, was a step which the Board could not 
take unless asked to do so by the Council. 

Of course, the latter body lost no time in eisee the | 
request, to which, however, they received the startling reply, 
that it had been discovered that the Act gave the Board no 
power to affiliate societies; it would, however, take the neces- 
sary steps to obtain such power. 

This led to the passing of a short Amending Act the 

following year, and affiliation between the two bodies was then 
duly effected. 

Another little matter, which resulted from the termina- 
tion of the union between the Society and the S.A. Institute, 
may be of interest as involving a principle. An early 
intimation was received from the Board, that after June 30, 
1884, the Society would be relieved of any further payment 
for the use of its room, except for the cost of gas consumed 
at its meetings and those of its branches; further, that in 
future it would receive no assistance in clerical work from 
the Board’s officers. 

It was about this time, also, that owing to the rapid 
growth of the School of Design, it became necessary for the 


Society to vacate its old room in the Institute and occupy a 


more commodious room in the new wing of the Museum. 
They remained there until 1891, when the School of Design, 
having removed to the Exhibition Building, the Society, with 


the consent of the Board, returned again to its old quarters 
in the Institute. 


(c) Establishment of Sections. 
As early in its history as 1858, a Committee was 


_ appointed to consider the expediency of dividing the Society 
into sections, each of which should be specially charged with 


the supervision of certain subjects. No less than ten such 
sections were proposed, and it was probably owing to their 
multiplicity that the scheme fell through. 

In 1883 the idea was revived, though from a different 
point of view. It was thought that there existed a need for 


a section of a popular nature, which would also serve as a 
recruiting ground from which the Society might increase its 


membership. Thus was established the Field Naturalists’ 
Section. It was intended for studiously disposed persons, of 
either sex, who wished to undertake the study of natural 


_ history from an elementary standpoint. 


Professor Tate, from whom the proposal had emanated, 


_ delivered an interesting lecture 1 in the Town Hall, explaining 
¥ 


i 


Be 


2 


644 


the objects of the Section. No scientific qualification was 
demanded from intending members. The Field Naturalists’ 
Section certainly met a public want, and has had a vigorous 
existence for about forty years. 

Its success led to the establishment of other Sections, 
such, for instance, as the Microscopical, the Malacological, 
etc., none of which, however, have survived. 


(d) The Society’s Library. 


It does not appear necessary to treat in detail the later 
developments and activities of the Society, for they are 
permanently. embodied in its Transactions and are compara- 
tively modern history. 

As the years passed the number of foreign exchanges was 
ever on the increase, until the library began to assume 
formidable proportions. As the room was small, and the 
shelving ludicrously inadequate, it became necessary to stack 
many of the books on top of each other, so that they were 
quite inaccessible for consultation by the Fellows. Frequent 
references are to be found in the annual reports in regard to 
this matter. In 1890 it is stated: ‘‘The Council is far from 
satisfied with the present conditions under which the books 
of the library have to be kept. It had been hoped that by 
this time arrangements might have been made to have them 
so placed in some portion of the Public Library that members | 
could have access to them at any time during the day. It 
feels that the present unsatisfactory condition cannot be 
allowed to continue, but that every effort must be made to 
place at the disposal of the Fellows the library in a more 
efficient way.’’ 

The next annual report shows, that during the year 
increased shelving accommodation had been provided, and that 
the books had been arranged in easily accessible positions. This 
report adds, that in order to make the library still more 
comprehensive and complete, the Council had put itself into 
communication with a number of American and European 
scientific societies, whose publications had been solicited in 
exchange for our own. 

Some ten years later it is announced, that ‘donations 
from scientific bodies have so largely increased of late, that 
the possibility of making the vast amount of material avail- 
able becomes a very urgent quéstion.”’ 

Accordingly a card catalogue was prepared, and this 
brought to light many breaks and irregularities in the sets 
of serial literature. A Committee was now appointed to 
inquire into the whole question of the library and its arrange- 
ment. This Committee came to the conclusion that the only 


a — 


645 


solution of their difficulties was the transfer of their books 
to the Public Library, because the Society had neither 
accommodation for the books nor a librarian to look after them, 

At this juncture the Government was approached with 
a view to securing better accommodation for the Royal and 
other local societies. The result was that additions were 
made to the northern end of the Institute Building, and fine 
premises were erected capable of. comfortably accommodating 
all affiliated Societies together with their respective libraries 
and property. It was completed and suitably furnished in 
1907, and the large western room on the ground floor, where 
we now hold our meetings, was allocated by the Board for 
the purposes of the Royal Society, a smaller room between 
this and the York Gate Library being apportioned for the 
common use of the two bodies. Under these greatly 
improved conditions, it was at last possible to introduce order, 
where chaos had formerly prevailed. 

As the Geographical Society occupied the adjacent room, 
it was at first thought that economy might be effected by the 
two Societies sharing the services of a single librarian, who 
should also act as their common Secretary. Unfortunately 


_ this scheme did not eventuate, and each Society subsequently 


appointed its own officer. 

The Society is under a debt of gratitude to Sir Joseph 
Verco for the great personal interest he has displayed in the 
reorganization of the library. Only those of us who remember 
the old order (or disorder ?) of things, can fully appreciate 


_ the nature of the change that has been effected during his 


Presidency. - 

In 1921, the following exchanges of publications were 
made with leaned societies in other countries :—United 
Kingdom, 27; Continental Europe, 66; Canada, 4; South 
Africa, 6; Sudan, 1; India and Ceylon, 6; United States, 
50; Mexico, 2; Brazil, 1; Uruguay, 1; Peru, 1; Argentina, 
i; Japan, > China, 1); Piao 1; Straits Settlements, 
1; Java, 2; Hawaiian Islands, ; Commonwealth of Australia 
and New Zealand, 54. oe 231. 


(e) Research and Endowment Fund. 
In 1903, the Society was incorporated, in order that it 
might acquire and hold property. 
The chief object of this was to enable it to establish 
an Endowment Fund, for the purpose of meeting special 


liabilities and also for the promotion of scientific research. 
Thanks to the generosity of Sir Joseph Vereo, the late 
_ Thomas Scarfe, and the late R. Barr Smith, each of whom 


donated the sum of £1,000, this fund has now been in opera- 


l _tion for some years. 


646 


5. ConcLusion. 


The total number of members on our roll is 102. Fully 
half of these have contributed papers which have been pub- 
lished in the Society’s Transactions, many of which must be 
regarded as very important additions to the literature of 
science. It is not my task, however, to particularize—it 
would, in fact, be invidious for me to do so. But there is 
one name, which I feel sure you would like me to mention— 
the honourable name of our senior Fellow, Walter Rutt, the 
connecting link between the old order and the new. Mr. 
Rutt was elected fifty-three years ago, when the Society was 
still in its callow youth, and during nearly the whole of that 
long period he has been a member of the Council, chiefly in 
the capacity of Hon. Secretary or Hon. Treasurer. He has 
also filled the office of Vice-President. He has accompanied 
the Society through all its vicissitudes of fortune, and is the 
authority to whom one naturally appeals for information on 
every important event in its history. Though he has contri- 
buted but few papers to its Transactions, yet in wealth of 
service he is probably its chief benefactor. 


APPENDIX I. 


List oF FounDATION MremMBERS ELECTED IN THE YEAR 1853. 


+Babbage, B. H. Hays, W. Bennett 
Babbage, Dugald Kay, Robert 

Bompas, Dr. J. C. Kingston, G. S. 
Brown, Dr. John Mann, Charles 

Clark, A. Sydney Martin, E. M. 

Clark, Francis Moore, Dr. R. W. 
*Clark, John Howard Moorhouse, Dr. Matthew 
+Davy, Dr. E. Mayo, Dr. George 
Davies, Dr. Chas. Nootnagel, H. 
Doswell, C. M. Quick, N. S. 
*Feinaigle, C. G. Shell, W,. Be 
Freeling, Capt. Arthur H. Stow, R. S. 

Garran, Dr. Andrew + Whitridge, W. W. R. 
Gilbert, W. B. - Williams, T. G. 
*Gosse, Dr. Wm. Wilson, C. A. 
Hamilton, Edward Wooldridge, Dr. H. 
Hamilton, G. E. Young, Sir H. E. Fox 
Hammond, Octavius *Young, John L. 


Hanson, R. D. 


“Attended the first preliminary meeting. 
tAttended the second preliminary meeting. 


647 


APPENDIX II. 


Officers of the Society. 


(a) *Past-PRESIDENTS. 


Sir H. E. Fox Young (2) Dr. H. T. Whittell (1) 
B. Herschel Babbage (1) Professor Horace Lamb (1) 
Sir R. G. MacDonnell (6) H. C. Mais (1) 


Sir Dominic Daly (7) Professor E. H. Rennie (6) 
Sir James Fergusson (5) Sir Edward Stirling (1) 
Sir Anthony Musgrave (4) Rev. Canon Blackburn (2) 
Sir William Jervois (1) Professor Walter Howchin (2) 
- Professor Ralph Tate (5) Dr. W. L. Cleland (3) 
_ Sir Samuel Way (2) Sir Joseph Verco (19) 
_ Sir Charles Todd (1) Dr. R. S. Rogers (1) 


(6) VicE-PRESIDENTS. 


_ Babbage, B. H. Moorhouse, Dr. M. 
we Freeling, Sir A. H. Gosse, Dr. Wm. 
i Davies, Dr. C. Light, W. H. 
| Farr, Rev. Canon G. H. Bruce, J. A. 
| Forster, Hon. Anthony Hanson, Sir R. D. 
| Todd, Sir Chas. Wilson, C. A. 
Maughan, Rev. Jas. Waterhouse, F. G. 
Schomburgh, Richd. M. Hanson, W. 
Hosking, James Smeaton, T. D. 
Ingleby, Rupert Bonney, Chas. 
_ Tate, Professor Ralph Chapple, Frederic 
_ Adamson, D. B. Whittell, Dr. H. T. 
Lamb, Professor Horace Stirling, Sir Edward 
Mais, H. C. Howchin, Professor Walter 
Mestayer, Richd. L. Dixon, Samuel 
Holtze, Maurice Rennie, Professor H. E. 
Blackburn, Rev. Canon T. Rutt, Walter 
Rogers, Dr. R. S. Pulleine, Dr. R. H. 
Verco, Sir Joseph Ashby, Edwin 


(c) REPRESENTATIVE GOVERNORS. 


Babbage, B. H., 1859-1860. 
Wyatt, Dr., 1860-1869. 
Todd, Sir Charles, 1869-1884. 
Whittell, Dr. H. T., 1884-1888. 
Tate, Prof. Ralph, 1888-1901. 
Howchin, Prof. Walter, 1901-1922. 
*The numbers enclosed in brackets indicate years of service. 


02 


648 
(d) *Hon. SECRETARIES. 
J. H. Clark (9) Walter Rutt (15) 
T. D. Smeaton (1) Dr. W. L. Cleland (15) 
James Hosking (3) W. C. Grasby (1) 
J. S. Lloyd (5) W. B. Poole (2) 
C. W. Babbage (6) G. G. Mayo (10) 
W. C. M. Finniss (3) Dr. R. H. Pulleine (3) 
(e) *Hon. TREASURERS. 
Charles Mann. (1) J. 8. Lloyd (2) 
Dr. Andrew Garran (2) Walter Rutt (23) 
A. Sydney Clark (8) W. B. Poole (12) 


John Howard Clark (11) B. 8S. Roach (1) 
T. D. Smeaton (8) 
(f) Epirors. 
Professor Ralph, Tate (16) Professor Walter Howchin (29) 


*The numbers enclosed in brackets indicate years of service. 


Bibliography. 
Adelaide Philosophical Society. 

*Annual Reports published by the Society. 1854-58; 
1865-73; 1858-9, in The Register, 27/7/59; 1860-1, 
in The Kegister,. 30/10/61; 1861-2, in MSS. 

Annual Reports (abstracts) included as appendices to the 
Annual Reports of S.A. Institute, 1863 to 1884. | 

*Correspondence with S.A. Institute (MSS. Ne 

*Exploring Exped. from Fowlers Bay into Interior, 
Correspondence relating to. 

*Incorporation with S.A. Institute, Papers relating to. 

*Laws of. Published January, 1854. 

*Minute-book (1853). 

*Papers read and deposited with the Society. 1853. 

*S.A. Acclimatization Society. Correspondence deena 
with Establishment of (MSS.). 

*Sections, Division into (MSS., etc.). 1859. 

*Title of ““Royal.’’ Correspondence relating to (MSS.). 

Educational Journ. of S.A. 15/8/57 and 15/9/57. Articles 
by Nathaniel Hailes on Early Societies. il 
Feinaigle, C. G. S.A. Almanac, 1851, p. 165; The Register, | 
23/9/63, p. 3; par. 7; Argus (Victoria), 18/3/80Re a) 
(obituary). = | 
Hack’s Expedition (1857), Parl. Pap. 156 and 189 of 1857-8. | 


*Charles Mann. Evidence before a Committee of the Adel. 
Library and Mechanics’ Inst. (1853). MSS. 


*In the Archives of the Public Library. 


: 


649 


_ Royal Society of S.A. 
*Correspondence with Public Library Board. 
i" Transactions of the Society, 1877-1921. 


The Advertiser. 
Vesting of Property of Incorporated Societies in the S.A. 
Institute, 26/7/61 (leading article). 


S.A. Institute. 

Appendices to Ann. Rep. (Adel. Phil. Soc.) for years 
ended 30/9/63 -30/9/84. 

Acts No. 16 of 1855-6; Amending No. 7 of 1856; Further 

‘amended (appointment of a 7th Governor) 1861; 

Consolidated, 1863; Ann. Reports, 1860-1 to 30/6/84 
(file of Pub. Lib.). 

Fittings; Correspondence relating to and payment for, 
Parl. Pap. No. 131 (28/8/60). 

Fittings; Petition to House of Assembly re (24/9/60), in 
file Pub. Lib. 

Minute-book, July, 1856-end 1858 (file of Pub. Lib.). 

Property of Incorporated Societies; Vesting of in S.A. 
Inst., The Advertiser (leader, 26/7/61). 

Inaugural Address, The Register, 30/1/61. 


_ S.A. Literary Association (named changed shortly after incep- 
tion to 8.A. Lit. and Sc. Assn.). 

Minute-book, 1834-5. 

Educational Journal, 15/8/57 (article by N. Hailes). 


The Register. 


Monthly and Ann. Meetings of Phil. Soc. reported 
regularly. 

Babbage’s paper on his proposed Exped., 3/2/58, p. 3. 

Feimaigle,; C..G., 23/9/63, :p..3, col. 7. 

S.A. Inst., site of, 3/5/59, p. 3, col. 6; 5/5/59, p. 3, 
col. 2. 

S.A. Inst., Inaugural Address, 30/1/61. 

Young, John L. pO ft plZ 5p. 10, col. 4. 


*The Southern Australian. 

Adelaide Mechanics’ Institute. 

“1838, June 30, p. 2, col. 2; Aug. 4, p. 2, col. 3; Aug. 18, 
piii2, col. 3.5 Sept. 1,.picb, col: 4; Oct! 27, <px3 
col. 3: Dec. 8, pi 2; col. 2. 

p1039, June 26, p. 2, col. 1, and p. 3, col. 1; Aug. 7, 
ped,.col. 3. 


*In the Archives of the Public Library. 


650 


PAPERS LAID ON THE TaBLE.—‘‘Insect Metamorphosis,” 
by O. W. Tires, M.Sc.; ‘‘Types of Species of Australian 
Polyplacophora now in the Museum of Natural History, | 
Paris,” by Epwin Asnsy, F.L.S., M.B.O.U.; ‘‘Additions 
to the Flora of South Australia, No. 20,” by J. M. Brack; | 
‘‘On a New Genus and Species of Australian Lycaeninae from 
the Roper River,’”’ by Norman B. TinpaLE (communicated by 
A. M. Lea, F.E.S.); ‘‘A Preliminary Note on the Fossil 
Woods from some Australian Brown Coal Deposits,’’ by Miss 
E. Dorotny Noses, B.Sc.; ‘‘On the Ecology of the Ooldea 
District,’”” by R. 8. Apamson, M.A., B.Se., and) 1) Guam, 
Ossorn, D:Sc.; “‘Designs on Rocks in the Burra District,”’ 
by Joun BIDDLE (communicated by A. G. Edquist); and 
‘‘Ecological Notes on South Australian Plants, Part I.,” by 
E. H. Isine. 

ELECTION OF OrriceRS.—Dr. Rogers declined renomina- 
tion as President. The following officers were elected for 
1922-23 :—President, R. H. Pulleine, M.B.; Vice-Presidents, 
R. 8. Rogers, M.A., M.D., and Sir Joseph C. Verco, M.D., 
F.R.C.8.; Hon. Treasurer, B.S. Roach; Alembers of Couneil, 
(vice Professors Chapman and Robertson, who retired by 
effluxion of time), Edgar R. Waite, F.L.S., and Professor 
T. G. B. Osborn, D.Sc.; Hon. Auditors, W. C. Hackett and 
H.. Whitbread. It was also decided to appoint an Hon. 
Assistant Secretary, and E. H. Ising was elected to the 
position. 


ANNUAL REPORT, 1921-22: 


Two lines of research have been followed this year with 
the assistance of grants from this Society. Mr. F. R. Marston, 
who is seeking to obtain from azine precipitate samples of 
the pure proteolytic enzymes, has forwarded to the Council a 
progress report of his work, which, as his investigations are 
incomplete, he desires them to be regarded as confidential. 
Prof. F. Wood Jones’ exploration of the Flora and Fauna 
of Nuyt’s Archipelago and the Investigator Group has 
resulted in the collection of valuable specimens, which are 
being dealt with by experts, and papers on the subject by 
Professors F. Wood Jones and T. G. B. Osborn, Dr. Chilton, 
and Mr. A. M. Lea will appear in our forthcoming volume 
of ‘“Transactions.’? The types of the new species obtained 
will be donated to the South Australian Museum. 

The largest contribution to this volume will be a paper 
on ‘‘Insect Metamorphosis’? by Mr. O. W. Tiegs, who also 


¥ 651 


furnishes a paper on “The Anatomy of the Voluntary 
_Muscles.’’ Further papers on the ‘‘Polyplacophora,”’ by Mr. 
Edwin Ashby, and the ‘‘Pouch Embryos of Marsupials,” by 
Prof. F. Wood Jones, will be printed, and various branches 
‘of Natural Science, Chemistry, and Ethnology will be dealt 
with by Professors Sir Edgeworth David, Walter Howchin, 
Sir Douglas Mawson, T. G. B. Osborn, and J. B. Cleland, 
Drs. E. O. Teale, R. S. Rogers, R. H. Pulleine, and A. 
_Jefferis Turner, Miss Nobes, and Messrs. J. M. Black, F. 
Chapman, A. G. Edquist, eA Berry, R. S. Adamson, 
_N. B. Tindale, and E. H. Ising. 


hi The exhibits at the evening meetings have been numerous 
| and interesting, including a series of views illustrating the 
Eucla Basin and its Water Supply, by Mr. L. Keith Ward. 


The Library is continually increasing through the ex- 
change of our publications for those of a growing number 
‘i of learned societies and other public bodies, but no definite 
reply has yet been received from the Government to the 
_ request made for additional shelving. 
| The Index to our publications for the years 1901-1920 is 
| in the printer’s hands and will be issued early. Its publica- 
| tion, which has been delayed by the great labour of revising 
| the proofs, will Jargely absorb the balance of our funds. 


I A suggestion having been made by the Queensland 
Branch of the Royal Geographical Society that a thorough 
exploration of the Great Barrier Reef should be made, Prof. 
_F. Wood Jones was appointed to represent this Society upon 
| a joint Committee to consider the proposal. 
i} We have to report the death during the year of three of 
“our Fellows—Dr. Sweetapple, of 15 years’ standing: Mr. W. 
| Ware, who was elected in 1878, and who served for many 
| years as Hon. Auditor; and Mr. F. R. Zietz, who since his 
| election in 1912 had taken an active part in the evening meet- 
ings by his papers, exhibits, and contribution to the 
discussions. 


The present membership of the Society comprises 
9 Honorary Fellows, 4 Corresponding Members, 87 Fellows, 
and 1 Associate. 
" R. S. Rocers, President. 


lie WaLteR Routt, Hon. Secretary. 


September 30, 1922. 


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654 


DONATIONS TO THE LIBRARY 


FOR THE YEAR ENDED SEPTEMBER 30, 1922. 


TRANSACTIONS, JOURNALS, REPORTS, ETC., 
presented by the respective governments, societies, and 
editors. 


AUSTRALIA. 


AUSTRALASIAN ANTARCTIC ExpEpiTiIon, 1911-14. ° Scientifie 
reports, ser. A, v. 3, pt. 1; ser. C, v. 5, pt. Gam 
6, pt. 2-3; 7, pt. 1-4; 8, pt. 1. Syd. 19te-28 
AUSTRALASIAN ASSOCIATION FOR THE ADVANCEMENT OF 
Scrence. Report, v.15. Melb. 1921. 
AUSTRALASIAN INSTITUTE OF MINING AND METALLURGY. 
Proc., no. 41-43. Melb. 1921. 
Australia. Bureau of Census and Statistics. Year-book 14. | 
Bureau of Meteorology. Rain-map, 1921. Melb. 
Fishertes, v. 5, pt. 2. Melb. 1921. 
Institute of Science and Industry. Bull. 22. Melb. 


SOUTH AUSTRALIA. 


Pusitic Liprary, Musrtum, anp Art GaLLeRyY oF S.A. 
Records of the S.A. Museum, v. 2, no. 2. Adel. 1922. 

Report, 1920-21. Adel. 1921. 

RovaL Greocrapuican Society (S.A. Br.). Proe., v. 21. 

Sourn AustRALia. Dept. of Mines. Government Geologist’s 
report, 1920. Adel. 1921. 

Review of mining operations, no. 34. 

Woods and Forests Dept. Report, 1920-21. 

SoutH AUSTRALIAN NATURALIST, v. 3. Adel. 1921-22. 

South AUSTRALIAN ORNITHOLOGIST, v. 6, pt. 4-7. 1921-22. 

Sourn AUSTRALIAN ScHoot oF Mines. Report, 1921. 


NEW SOUTH WALES. 


AUSTRALIAN Museum. Magazine, v. 1, no. 3-5. 
—_+—. ‘Records, y,-13, mio, 5. 8 Syd. glo22, 
aa Report, 1920-21. Syd. a 
Linnean Society oF N.S.W. Proc., v. 46; 47, pt. 1-2. a 
Marpen, J. H. Critical revision of the genus Eucalyptus, | 
pt. 49-56. Syd. 1921-22. | 
Forest flora of N.S.W., v. 7, pt. 5, 8-10. 1921-22. | 
New Soutn Watss. Botanic Gardens. Report, 1920. — 
Dept. of Agriculture. Gazette, v. 32, no. 10-12. 
— Science bull., no. 19, 22. Syd. 1921-22. 
Dept. of Mines. Annual report, 1921. 


—— 


Sr 8 


; 


655 


New Sourn Wates. Dept. of Mines. Geological memoirs, 


no. 8, 1922. 
: Public Inbrary. Report, 1921. Syd. 
Rovyat Socrery or N.S.W. Journ., v. 54. Syd. 1920. 
Sypney University. Calendar, 1922. 


QUEENSLAND. 


€ 


QUEENSLAND. Dept. of Agriculture. Journ., v. 16, pt. 4, to 


we te. pl: 3: 
QUEENSLAND Museum. Mem., v. 7, pt. 3. Brisb. 1921. 


- Roya Society oF QUEENSLAND. Proc., v. 33, 1921. Brisb. 


TASMANIA. 
Roya Society of TasMANIA. Proc., 1921. »Hobart. 
Tasmania. Dept. of Mines. Underground water-supply paper, 
no. 2. Hobart. 1922. 


VICTORIA. 
Royat Society or Victoria. Proc., v. 34, pt. 1-2. 1921. 
ScIENTIFIC AUSTRALIAN, v. 27, no. 7-12; 28, no. 1-6. Melb. 
Victoria. Dept. of Agriculture. Journ., v. 20, pt. 3-9. 
Geological Survey. Bull., no. 45. 1922. 
Records, v. 4, pt. 3. Melb. 1921. 
National Museum. Ethnological guide. 1922. 
VicToRIAN NATURALIST, v. 38, no. 6, to v. 39, no. 5. Melb. 


WESTERN AUSTRALIA. 
Royau Society or W.A. Journ., v. 7. Perth. 1921. 


ENGLAND. 


British Museum (Nat. Hist.). Economic ser., no. 12. 


Report on cetacea stranded on British coasts, 7. 


-CamBripce PuinosopHican Society. Proc., v. 20, pt. 4; 


21, pt. 1-2. 

Trans., v. 22, no. 23-25. Camb. 1922. 

CaMBRIDGE UNIvERSITY. Solar Physics. Report, 1920-21. 
ConcHoLocicaL Society. Journ., v. 16, no. 7-9. 1921-22. 
EntTomoLoeican Society. Trans., 1921, pt. 1-2, 5. Lond. 
GrEoLoGicaL Society. Journ., v. 77, pt. 2-4; 78, pt. 1-2. 
Hitt Museum. Bull., v.1, no. 1. Witley. 1921. 
ImpeRiaAL Bureau oF Entomotocy. Review of applied entom- 
ology, ser. A and B, v. 9, pt. 8-11; 10, pt. 1-7. 


| Imperiau Institute. Bull., v. 19; 20, no. 1. 1921-22. 
Linnean Society. Journ., botany, no. 303-5. 1921-22. 


Journ., zoology, n. 230-31. Lond. 1922. 
: List and proc., 1920-21. Lond. 
Trans., zoology, v. 18, pt. 1. Lond. 1922. 


‘Liverroot Bioroeicat Society. Trans), vi. 35s.) \1921 . 


! 


cmd 


656 


MANCHESTER LITERARY AND PHILOSOPHICAL SocieTY. Mem. 
and proc., v. 64, pt. 2; 63, pt. 1. 1921. 

Royat Boranic GARDENS, Kew. Bull., 1921. Lond. 

Hooker’s icones plantarum, v. 1, pt. 4. 1922. | 

RoyaL GEOGRAPHICAL Society. Journ., v. 58, no. 3, to 60, 
no. 2. 

Royat Microscopicau Society. Journ., 1921; 1922, pt. 1-2. 

Royaut Society. Proc., A 701-12, B 647-56. 1921-22. 


— 


Year-book. 1922. Lond. ; 
UnitepD Empire, v. 12, no. 9, to 13, no. 8. 1921-22. 
SCOTLAND. 
Royat Society oF EpinpureH. Proc., v. 41, pt. 2, to 42, 
pie a ae | ; 
Trans. ,' v.52) pt.i4:" Edin. 1927, 
IRELAND. 


Royat Dusuin Society. Scientific proc., v. 16, no. 14-34. 


ARGENTINE. 


AcADEMIA NACIONAL DE CIENCIAS EN Corpopa. Bull., t. 24, 
pt. 3-4; 25, pt. 1-3; 26, pt. 1. 1921-22. 


AUSTRIA. 


AKADEMIE DER WISSENSCHAFTEN. Anz., 1921. Wien. 
Mitteilungen, no. 47-57. 1914-19. 

Sitz., Bd. 123-130. Wien. 1914-21. 

GEOLOGISCHE STAATSANSTALT. Verh., 1921. Wien. 
NATURHISTORISCHEN HormusEeums. Ann., Bd. 34-35. ' 
Zoou.-Bot. GESELLSCHAFT IN WIEN. Bd. 70-71. 1921-22. 


BELGIUM. 


ACADEMIE ROYALE DE BELGIQuE. Ann., 1922. Brux. 
L’ académie depuis sa fondation, 1772-1922. 
— — Classe des Sciences. Bull., 1921, no. 4, to 1922, no. 4. 
Mem. in 4°, ser. 2, t. 4, fasc. 6-7. | 
Mem. in 8°, ser. 2, t. 6, f. 4, 7-15. 
Institut RoyvaL MéréoroLocique. Ann., t. 1, f. 1. 19195 || 
Muss&e Roya Dd’ Historre NaturELLE. Ann., t. 1-13, text | 
and plates. Brux. 1876-96. | 
Mem., t. 8, fasc. 4. Brux. 1921. , Ei 
OBSERVATOIRE RoyaL DE Bencigue. Annuaire, 1923. | 
Annales, ser. 3, t. 1, fasc. 1. Brux. 1921. . ol 
SociETs ROYALE DE Botanique. Bull., t. 50, 54. 1921. eo 
SociiT& RoyaLeE DES ScIENCES DE LifcE. Mem., s. 3, t. ll; |} 
fy . 
SocriTE RoyaLe ZooLoGiquE ET Mauacoxtogiague. Ann. 51-2. | 


+ 
& 


> 


——————E 


caeetiaiimtoee. 


657 


BRAZIL. 


Instituto OswaLpo Cruz. Mem., t. 13, fasc. 1. 1921. 
OBSERVATORIO NACIONAL DO R10 DE JANEIRO. Anno 38. 


CANADA. 


Canapa. Geological Survey. Mem. 124-128. Ottawa.’ 
Museum bull. 36; and various publications. 
Mines. Bull. 33-34; and various publications. 
Canapian Arctic ExPepDiTIon, 1913-18. Report, v. 12. 
CANADIAN INSTITUTE. Trans., v. 13, pt. 2; 14, pt. 1. 
Nova Scotian Institute oF Science. Proc., v. 15, pt. 1. 
Roya Society oF CanaDA. Proc.; v. 15, 1921. Ottawa. 


CEYLON. 
Cotomso Museum. Spolia Zeylanica, pt. 45. 1922. 


CHINA. : 
Roya Asiatic Society, Nru.-Cuina Br. Journ., v. 52. . 


DENMARK. 


ConsEIL PERMANENT INTERNATIONAL POUR L’ KXPLORATION DE 
LA Mer. Bull. 1905-18. Cpng. 

Dansk Botanisk Forenine. Arkiv., Bd. 2, no. 6. Cpng. 

Dansk NATURHISTORISK FoRENING. Vid.-med., Bd. 72. 

K. DANSKE VIDENSKABERNES SELSKAB. Biol.-med., Iil., 2. 

KOBENHAVN UNIVERSITETS ZOOL. Museum. Bull. 19-25. 


FRANCE. 
SOcIETE DES SCIENCES NATURELLES DE L’ OUEST DE FRANCE. 
Bull., t. 6. Nantes. 1920. 
SoOcIETE ENTOMOLOGIQUE DE FRANCE. Ann., 90, pt. 1-2. 
Ball, 2921) no. 13-21-1922. no. 1-12.° “Par. 
Societe LINNEENNE DE BorpEaux. Actes, v. 71-72. 1919-20. 
SocigETE LINNEENNE DE NORMANDIE. Bull., ser. 6, v. 7-10; 
ser. 7, v. 1-3. Caen.” 1914-21. 


GERMANY. 


BERLINER GESELLSCHAFT FUR ANTHROPOLOGIE. Zeits. fur 
Ethnologie, 1920-21, H. 4-6. 1921. 
DevutscHE ENntTomMoLociscHE Museum. Entom. Mitteil., Bd. 

fet it mne, t-4.) Berl, ) 1912-22. 

Freppe, F. Repertorium, Bd. 17, no. 13-30. Berl. 1921. 
GESELLSCHAFT DER WISSENSCHAFTEN ZU GOTTINGEN. Nach., 
Gesch. Mitteil., 1921; Math.-Phys., 1921, H. 2. 
GESELLSCHAFT FUR ERDKUNDE. Zeits., 1914, no. 7-9; 1921,- 
no. 3-10. Berl. 1914-21. : . 


658 


NASSAUISCHE VEREIN FUR NATURKUNDE. Jahrg. 73. Wiesb. 
NATURHISTORISCHE GESELLSCHAFT, Nwtrnsperc. Abh., 21, 
tite 

Jahr, 1919-20; 

NatTuRHISTORIScCHE Museum, Hamsure. Mitteil., v. 30, 38. 
PREUSSISCHE AKAD. DER WISSENSCHAFTEN. Sitz., 1914-21. 
Phal.-hist. Klasse. Sitz., 1922, no. 1-14. Berl. 
Phys.-math. Klasse. Sitz., 1922, no. 1-12. 
STADTISCHE MusEuUM FUR VOLKERKUNDE, LEIpzic. Jahr. 6-8. 
Veroff, H. 4-5. 1912-14. 


—- 


ee 


HAWAIIAN ISLANDS. 


BERNICE Pauanui BrsHop Museum. Mem., v. 8, no. 1-4. 
Occasional papers, v. 7; 8, no. 1-3, 5. 
Hawaiian ENtToMOLOGICAL Society. Proc., v. 4, no. 3. 


HOLLAND. 
Muster Teyter. Archives, ser. 3, v. 3-4. Haarlem. 1917-19. 
Risk’s Hersartum. Mededeel., no. 38-41. Leiden. 


HUNGARY. 
Museum Narrionatis. Ann., v. 17-18. Budapest. 1919-21. 


INDIA. 

. Agricultural Research Inst., Pusa. Bull. 101-113, 115. 
Report, 1920-21. Cale. 
Board of Scientific Advice. Report, 1920-21. 
Dent. of Agriculture. Botanical mem. 11, no. 4-5, 7. 
Chemical mem. 6, no. 1-2, 4-5. Cale. 
Entomological mem. 7, no, 6. 
——— Report of 3rd. Entomological Meeting, 1921. 
Review of agricultural operations, 1920-21. 
Geological Survey. Mem., v. 48. Cale. 1922. 
———— Palaeontologica Indica, n.s., v. 6, no. 2. 
Records, v. 53, pt. 2-3. Cale. 1921-22. 
Mapras. fisheries. Bull. 12-13. Reports 3-6, 1921; 1-2, 

1922. 
Royat Asratic Society, Bompay Br. Journ., v. 25, no. 3. 


| 
Z 
o 
4 
> 


TLL 


Maurieuia, anno. 29, fasc. 1-6. Catania. 1921. 

SocrETA DiI ScrENzZE NaTURALI ED EconomicHE. Giornale, 
v. 26, 28. Palermo. 1908-11. 

SocrETA IrTaLIaNa DI ScIENZE Naturaui. Atti, v. 60-61, f. 1. 

SocretaA Toscana DI ScIENZE Naturati. Mem., v. 26, 34. 

Processi verbali, v. 30. Pisa. 1921. 

Torino, R. Universita pi. Museum bull. 34-36. 


659 


JAPAN. 


Japan. Imperial Earthquake Investigation Committee. Bull., 
ve tOe no: i) Pokyo: | 1922. 

: Seismological notes, no. 1-2. 1921-22. 

Kyoto Imperiat Univ. College of Science. Mem. 5, no. 5-6. 

Nationa ResEarcH Councit. Japanese journal of astronomy 
and geophysics, v. 1, no. 1. Tokyo. 1922. 

; Japanese journal of geology, v. 1, no. 1. 

Proe., ue. 1. Tokyo... 1922. 

ToHnoku Imperial Univ. Science reports, Ist. s., v. 10, no. 3, 
ivy) ono..2; 2nd. s., y. 6, no. 1. Sendaz. 

Technology reports, v. 2, no. 2-4. 1921-22. 

TéHoKU MATHEMATICAL JOURNAL, v. 19-20; 21, no. 1-2. 

Toxyo ImprertaL Univ. College of Science. Journ., v. 41, 
gre) (-1b 42 art. 23°43). art. 7-8.; 1921. 


é; JAVA. 
NATUURKUNDIGE VEREENIGING IN NeEDERL.-INDIE. Tijds., 
Deel 81, pt. 2-3; 82, pt. 1-2. Weltevreden. 
NeEpDERL. Oost-InDIE Mynwezen. Jaarb., 1918, pt. 2. 


MEXICO. 


SocreDAD CIENTIFICA ‘‘ANTONIO ALZATE.” Mem., t. 35, 
no. 5-12; 39; 40, no. 1-6. Mexico. 1921-22. 


NEW ZEALAND. 
AUCKLAND INSTITUTE AND Museum. Report, 1921-22. 
CanTERBURY MusEeum. Records, v. 2, no. 2. Christchurch. 
New ZEALAND. Board of Science and Art. Bull., no. 2. 
N.Z. journal of science and technology, v. 4, 
ao, 9-07 5, to. 1-3. -Well. 1921-22. 
Dept. of Mines. Palaeontological bull., no. 9. 
Dominion Laboratory. Report, no. 54. Well. 1921. 
_Domimon Museum. Report, 1918-21. Well. 
Geological Survey. Annual report, no. 15-16. 
Ball, ue. 25.- Well.’ 1921. 
New ZeEaLanD InstituTe. Trans., v.53. Well. 1921. 


NORWAY. 
Bercens Museums. Aarsberetning, 1919-21. 
Aarbog, 1918-19, H. 2; 1919-20; 1921, H. 4-3. 
K. Norske VIDENSKABERS SELsKABS. Skr., 1918-20. 
STAVANGER Museum. Aarshefte, 1920-21. 


660 


PERU. 
ASOCIACION PERUANA PARA EL PROGRESO DE LA CIENCIA. 
Archivos, tom. 1, fase. 1. Lima. 192): 
CUERPO DE INGENIEROS DE Minas. Anales, t. 1-2. 
Bull. no. 55, 101-1038) Tania. 1807=2 10 


PHILIPPINE ISLANDS. 


BUREAU OF ScrENCE. Journ., v. 18, no. 6, to 20, no. 5. 
Mineral resources, 1919-20. Manila. 1922. 


SPAIN. 
InsTITUTO GENERAL Y TECNICO, VALENCIA. An., v. 7. 
REAL ACADEMIA DE CrENcTIAS y ARTES. Bull., os 4, no, D. 
Mem., v. 16, no. 6-11. Barcelona. 1920- 21. 


STRAITS SETTLEMENTS. 
Royau Asiatic Society, Straits Br. Journ., no. 84-85. 


SWEDEN. 


ENTOMOLOGISKA FORENINGEN 1 StocKHoLM, Tidsk., arg. 42. 

GEOLOGISKA FORENINGEN I StockHoutm. Forh., Bd. 44, H. 1-4. 

Reeia Socretas ScrentiaRuM Upsauiensis. Nova acta, 
ser, 4,’ v.50) dase. 44 Uipsalas toate 


SWITZERLAND. 
Institut NaTionaL GENEvoiIs. Bull., t. 44. 1921.. 
NATURFORSCHENDE GESELLSCHAFT IN ZUrRicH. Viert., 1921. 
SoclETE DE PHYSIQUE ET D’ HisrorrE NATURELLE. Comte 
rendu des séances, v. 38, no. 2-3; 39, no. 1-2. 
Mém., v. 39, fasc. 6-7. Geneva. 1921. 
SociETE VAUDOISE DES SciENCES Nat. Bull. 201-3., 


UNION OF SOUTH AFRICA. 


DurBan Museum. Annals, v. 3, pt. 2. 1921. 

GroLtocicaL Society ofr 8.A. Trans., 1921. Johannesb. 

Roya Society oF §.A. Trans., v. 10, pt. 1-2. Cape Town. 

S.A. ASSOUIATION FOR THE ADVANCEMENT OF SCIENCE. 
Journ. of science, v. 18, no. 1-2. Cape Town. 1921. 

S.A. Muservum. Annals, v. 18, pt. 3-4. 1921. 

Report, 1921. Cape Town. 


UNITED STATES. } 
ACADEMY oF NaTuRAL Sciences, Puoitap. Ann. rep., 1920. 
Proc., ¥. 112, pt. a3 73, pt. 1-3. 1920. 
ACADEMY OF ‘ScrENcE, Sr. Louts. Tr., v. 22, no. 4°637am 
BO. tT wo? 


\ 


661 


AMERICAN ACADEMY OF ARTS AND SCIENCES. Mem. 14, no. 3. 

Proc., v. 56, no. 1-4, 9-11; 57, no. 1-10. Bost. 

AMERICAN CHEMICAL Society. Journ., v. 40, no. 4-10; 43, 
no. 6-12; 44, no. 1-7. Easton, Pa. 1918-22. 

AMERICAN GEOGRAPHICAL Society. Review, v. 11, no. 4, to 
£2 no. 3; 

AMERICAN INSTITUTE OF MiniInG ENGINEERS. Tr., v. 64-66. 

AmeERIcAN Microscopical Society. Tr., v. 40-41, no. 2. 
AMERICAN Museum oF Natura History. American Museum 

novitates, no. 1-42. N.Y. 1921-22. 

Anthropological papers, v. 16, pt. 6-7; 20, pt. 2; 23, 
pt. 4-5; 25, pt. 2; 26, pt. 2; 27. 
Bull., v. 42-44; 46, art. 1. N.Y. 1921-22. 

Guide leaflets, no. 51-54. 1921-22. 
Mem., n.s., v. 3, pt. 2-3. 1920-21. 

‘Natural History,’’ v. 21, no. 3-6; 22, no. 2-3. 

frepory, 1920." N.Y. "1922: 

AMERICAN PuHILOSoPHICcAL Society. Proc., v. 60. 

ARNOLD ARBORETUM. Journ., v. 2, no. 4. Camb., Mass. 

Boston Society oF NaturaL History. Mem., v. 8, no. 3. 

Proc., v. 35, no. 4-6. Bost. 1917-20. 

BrRookLtyn INSTITUTE oF ARTS AND ScIENcES. Museum 
quarterly, v. 6, no. 4; 8; 9, no. 1-2. 1919-22. 

BuFFALo Society oF NatTuRAL ScIENcCES. Bull. 13, no. 2. 

CALIFORNIA ACADEMY OF SCIENCES. Proc., v. 9, no. 14-15; 
ine. 10 cht no. V 1-17) San’ Fran. 1920-21. 

CALIFORNIA STATE Minine Bureau. Bull., no. 75, 82-83, 86, 
88-90. Sacramento. 1920-21. 

Oil-fields summary, v. 5, no. 6-11; 6-7. 

Report, now8,17;)v. 18, no. 1-6. 1921-22. 

CaLiFoRNiaA UNivEeRsity. Mem., v. 5. Berkeley. 1921. 

Publications in agriculture, v. 4, no. 8. 

Archaeology and ethnology, v. 12, no. 2-7. 

Botany, v. 5, no. 9-11; 7, no. 1-4. 1916-17. 

Geology, v. 10, no. 2-10. Berkeley. 1916-17. 

Lomo: to, mowth: 13; no 13; 15) ne: 2-316; 
Hoo reo wh neta eO 19 no. 6:20, no. fs 2s 

ConnEcTIcuT ACADEMY OF ARTS AND ScIENCES. Trans., v. 15, 
pp. 93-408. New Haven. 1921-22. 

Connecticut Gro.ocicaL Survey.. Bull. 5-6. Hartford. 

CoRNELL University. Agricultural Experiment Station. 
Bull. 26-37, 50-61, 392-99, 403-7. 

Mem., 12-13, 19, 21, 24-25, 27, 34-52. 

Report, 1888-1913, 1919-20. Ithaca, N.Y. 

Denison ScientiFic Association. Bull., v. 17, art. 1-7; 
18, art. 4-7; 19, art. 9-16. Granville, O. 

Fretp Museum. Zool. ser., v. 14, no. 1. Chic. 


II 


it 


662 


FRANKLIN INSTITUTE. Journ., v. 192-4, no. 2. Philad. 
Harvarp CoLtLtEGE. Museum. Ann. report, 1920-21. 

Bull., v. 65, no. 1-4. Camb., Mass. 
ILLINOIS UNIVERSITY. Biological monog., v. 6, no. 1-4. 
Agr. Expervment Station. Bull. 116, 178, 180, 186-7. 
Inp1aNA ACADEMY OF SCIENCE. Proc., 1919-20. Indianap. 
Jouns Hopkins UNIvERsItTy. Cire., 1920, no. 2-5. Balt. 
Studies in historical and political science, v. 38-39. 
Kansas University. Bull., humanistic, v. 1-2. 

Science bull., v. 13, no. 1-9. Lawrence. 1920. 
_LELanp STANFORD UNIVERSITY. Univ. ser., no. 36-43. 
Minnesota UNIVERSITY. Current problems, no. 13. 

Agr. Experiment Station. Bull. 190-93, 195-97. 
Missouri Botanic GARDEN. Ann., v. 1-7; 8, no. 2-3. 
NationaL ACADEMY OF SCIENCES. Proc., v. 3, no. 3; 6, no. 2; 
7, no. 3-10, 12: 8).me., 1-7. Wash: ot 9ijozae 

National Research Council. Bull., v. 2, pt. 3, no. 11. 
New York Pusiic Liprary. Bull. 25, no. 8, to 26, no. 7. 
New York State Museum. Report, 1916-18. 

New York Zoonocicar Society. Zoologica, v. 2, no. 12-13; 
3, no. 1-13. ONY, “1921. 

Zoopathologica, v. 1, no. 6. 1921. 

Nortu Carouina. Geological Survey. Cire. 1-3. 1922. 
Economic papers, no. 51-52. Raleigh. 1921. 
OBERLIN CoLLEGE. Wilson bull., v. 28, no. 4. 1916. 
Oxni1o University. Bull., v. 25, no. 26; 26, no. S,1o: 
Ohio. journal of science, v. 22, no. 1-5. 

OxiaHoma. Geological Survey. Bull. 26-27. Norman. 
SMITHSONIAN INSTITUTION. Annual report, 1917-20. 
Bureau of American Ethnology. Bull. 72, 74. 
Report 35-36, 1913-15. Wash. 

STANFORD UNIVERSITY. Biol. ser., v. 1, no. 2-4. 

Math; ser.) vy. 1), no. 1.) 920. 

TENNESSEE. Geological Survey. Bull. 25-26. Nashv. 
Unitep States. Coast and Geodetic Survey. Rep., 1921 
Results of observations, 1917-18. 

Special publications, various. 

Dept. of Agriculture. 10 bull. of dept., and 5 farmers’ 
bull. Wash. 
Entomological ser., no. 16, 17, 20. 

——— ———_— Experiment station record, v. 44; 45, no. 1-6, 

8-9; 46, no. 1-7. Wash. 

— Journ. of agricultural research, v. 13, no. 1; 

19, no: 7, 12; 21, no.. 8-12; 22, no. 1-9: 19a | 

North American fauna, no. 45. 

—___ —____ Year-book, 1920. Wash. 1921. 
| 
| 
: 


~ Pe Nem ee 


re 


663 


Unitep Sratres. (Geological Survey. Ann. rep., 42; also 
many bull., mineral resources, and water-supply papers. 

Geologie folios 211-213, and many sheets. 

Prof. papers 121, 123, 128, 129-1. 

— Library of Congress. Rep., 1920-21. Wash. 

—— National Museum. Ann. rep., 1921-22. Wash. 

ole 825 vey pb. 25° LOO we Ae pb. Ov. 74; 


— 


112-119. 
—— ~ Contrib. from Nat. Herbarium, v. 20, pt. 
10-12; 22, pt. 4-6; 24, pt, 1. Wash. 1921-22. 
Proc. Vv. 0l-oo.). Wash. “1921. 
WaGner FREE INSTITUTE OF ScIENCE. An., 1921-22. 
Mipans.. Voadeipes 2a etbalad. 1921. 
WASHINGTON UNIVERSITY, St. Louis. Humanistic ser., 
Veet oxen ls C917-20. 
Selentine cer) v.39, pt... no, 2); 4, pt. 2. imo. ; 
Gop. 2,9; pt. [-2.. Sb. Louis, Mo.» 1916-21. 


URUGUAY. 
Museo Nactonau. An., s. 2, v. 1, pt. 4. Montevideo. 


664 


LIST OF FELLOWS, MEMBERS, ETc. 


AS EXISTING ON 


SEPTEMBER 30, 1922. 


Those marked with an asterisk have contributed papers pub- 
lished in the Society’s Transactions. 


Any change in address should be notified to the Secretary. 


Nore.—The publications of the Society will not be sent to 
those whose subscriptions are in arrears. 


ee : Honorary FELLOWS. 


1910. *Brace, Sir W. H., K.B.E., M.A., D.Sc., F.R.S., Pro- 
fessor of Physics, University College, London (Fellow 
1886). 

1893. *Cossmann, M., 110, Faubourg Poissonnieére, Paris. 

1897. *Davip, Sir T. W. "EDGEWORTH, K.B.E. C.M.G., D.S.0O., 
Bell D.Se.. Res... F.G:S., Professor of ’ Geology, 
University of Sydney. 

1905. Guitt, THomas, C.M.G., I1.8.0., Glen Osmond. 

1905. ee CHas. is Assistant Curator, Australian Museum, 

ney 

1892. Mebane ue oie I.S.0., F.R.S., F.L.S., Director Botanic 
Gardens, Sydney, New. South Wales. 

1898. *Meryricr, E. aD , i F.Z.S., Tohrnhanger, Marl- 
borough, Wilts, eae: 

1894. *Witson, J. T., M_D., Ch.M., Professor of Anatomy, 
Cambridge University, England. 

1912. *Trrrrr, J. G. O., F.L.S., Elizabeth Street, Norwood 
(Corresponding Member 1878, Fellow 1886). 


CoRRESPONDING MEMBERS. 
1918. *Carter, H. J., B.A., Wahroonga, New South Wales. 
1909. *JoHncocrk, C. F., “Clare. 
1905. THomson, G. M., "ELS. Dunedin, New Zealand. 
1908. *WooLNoueH, WALTER Guorae, D. Se., F.G.S. (Fellow 1902). 


FELLOWS. 


1895. *Asupy, Epwin, F.L.S., M.B.O.U., Blackwood. 

1917. Baruey, J. F., Director Botanic Garden, Adelaide. 

1902. *Baxerr, W. H., ¥.L.S., King’s Park. 

1921. Binxs, Menviiee, M.B., B.S., F.R.C.S., Hospital, Broken 

il 

1902. *Buacxk, J. McConnett, 82, Brougham Place, North 
Adelaide. 

1912. *Broveuton, A. C., Young Street, Parkside. 

1911. Brown, Evear J., M.B., D.Ph., 3, North Terrace. 

1883. *Brown, By. Vin 286, Ward Street, North Adelaide. 

1916. * Bri, Lionet B., D.V.Sc. MS Laboratory, Adelaide Hospital. 

1921. Burton, R. J., Fuller Street, Walkerville. 

1922. Campsety, T. D. BaD pc Adelaide Hospital, 

1907. *CHAPMAN, R. W., M.A.  B.C.E., F.R.A.S., Professor of 
Engineering ‘and Mechanics, "Univ ersity of Adelaide. 


1922. 
1904. 
1895. 


1907. 


1916. 


1887. 


1915. 


1921. 
1911. 
1902. 
1918. 
1917. 


1914. 


TOTS: 
1904. 


1880. 
1910. 


1922. 
1904. 


LOUG. 
1922. 


1922. 
1916. 


1896. 
1883. 


1918. 
1912. 


1923. 
1918. 


1910. 
1921. 
1920. 
1918. 
1915. 


1897: 
1884. 


1922. 
1922. 
1888. 


665 


CuartEs, Aubert G., 88, Spring Street, Queenstown. 

CHRISTIE, W., 49, Rundle Street, Adelaide. 

*CLELAND, J OHN B., M.D., Professor of Pathology, Univer- 
sity "of Adelaide. 

*Cooxe, W. T., D.Sc., Lecturer, University of Adelaide. 

DARLING, H. Git Franklin Street, Adelaide. 

*DIxon, SAMUEL, Bath Street, New Glenelg. 

Dopp, Aan P., Prickly Pear Laboratory, Sherwood, 
Brisbane. 

Durron, G. H., B.Sc., F.G.S., University of Adelaide. 

Dutton, H. H., B.A. (Oxon.), ” Anlaby, 

*Epquist, A. G 2nd Avenue, Sefton Park. 

*EKuston, A. H., B.ES., Lefevre Terrace, North Adelaide. 

* KENNER, Cuas. A. E., cub) Sem ue Case: Education Depart- 
ment, area 

FERGuson, . W., M.B., Ch.M., Gordon Road, Roseville, 


cee 

GuastonsuRY, O., Adelaide Cement Co., Brookman 
Buildings. 

Gorpon, Davin, c/o D. & W. Murray, Gawler Place, ~ 
Adelaide. 


*GoypEeR, Groraz, A.M., F.C.S., Gawler Place, Adelaide. 

*GRANT, "KERR, M.Sc. be Professor of Physics, University of 
Adelaide. 

Grant? ROE. T., M-B:, BS., M.R.C.P., University of 
Adelaide. 

GrirFitH, H., Brighton. 

HACKETT, W. ise 35, Dequetteville Terrace, Kent Town. 

Hate, H. M., Molesworth Street, North Adelaide. 

“Ham, W ILETAM, F.R.E.S. University eae Adelaide. 

Hancock, : Lipson, A.M.I.C.E. euvis I1.M.M., M.Am.I.M.E., 
Kennedya, Wallaroo Mines. 

Hawker, HE. W., F.C.S., East Bungaree, Clare. 

*Howcuin, PROFESSOR WALTER, F.G.S., ‘Stonycroft,’’ 
Goodwood East. 

Isine, Ernest H., Loco. Department, Islington. 

Jack, RK. L., B.E., Assistant Government Geologist, 
‘Adelaide. 

James, THomas, M.R.C.S., 9, Watson Avenue, Rose Park. 

JENNISON, Rev. J. OC., Crocodile Islands, Northern 
Territory. 

*Jounson, EH. A., M.D., M.R.C.S., 295, Pirie Street, 
Adelaide. 

*Jounston, Proressor T. Harvey, M.A., D.Sec., Univer- 
sity of Brisbane. 

JONES, > Weop) iy M.B.. i B.8:, . M.B.C.S.5) GR. CrP. 
D.Sc., Professor of Anatomy, University of Adelaide. 

Kimser, W. J., Gaza. 

*Laurif£, D. F., Agricultural Department, Victoria Square. 

*Lea, A. M., F.E.S., South Australian Museum, Adelaide. 

Lenpon, A. A., M.D. (Lond.), M.R.C.S., Lecturer in 
Obstetrics, ” University of Adelaide, and Hon. 
Physician, Children’s Hospital, North Adelaide. 

LENDON, ALAN’ H., North Terrace. 

Lenpon, Guy A., 'M.B., B. S., M.R.C.P., North Terrace. 

*LOWER, OswaLp B., ¥.Z.S.., F.E.S., Broken Hill, New 
South Wales. 


666 


Mapiean, C. T., B.A., B.Se., University of Adelaide. 

MarHews, G. M., F.R.S.E., F.L.S., F.Z.S., Foulis Court, 
Fair Oak, Hants, England. 

*Mawson, Sir Doveras, D.Sc., B.E., Professor of Geology, 
University of Adelaide. 

re Hersert, LL.B., Brookman Buildings, Grenfell 
treet. 

Mayo, Herren M,. M.B., B.S., 47, Melbourne Street, 
North Adelaide. 

McGitp, Joun Nett, Napier Terrace, King’s Park. 

MELROSE, ROBERT THOMSON, Mount Pleasant. 

*Morean, Aly MEME Ch, B:, 46, North Terrace, Adelaide, 

Movwu.LpEN, OwEN Vi. ‘M.B., B. S., Broken Hill. 

*“NOBES, Epiri D., B.Sc. - University of Adelaide. 

*OsBorRN, Ge oe D.Sc. ., Professor of Botany, University 
of Adelaide. 

Pootz, W. B., 6, Rose Street, Prospect. 

PopE, WILLIAM, Eagle Chambers, Pirie Street. 

*PULLEINE, R. H., M.B., 3, North Terrace, Adelaide. 

Ray, WIL1iamM, M. B.; B. Se., Victoria Square, Adelaide. 

*RENNIE, EDWARD H., M.A., D.Sc. (Lond.), V.C.S., Pro- 
fessor of Chemistry, University of Adelaide. 

Roacu, B. §., Education Department, Flinders Street, 
Adelaide. 

*RoBERTSON, Proressor T. B., University of Adelaide. 

*Rogers, R. S., M.A., M.D., Hutt Street, Adelaide. 

*Rorr, WALTER, C.E., College Park, Adelaide. 

SELWaAy, Weck. , Treasury, Adelaide. 

*SAMUEL, Guorrrey, B.Sc., University of Adelaide. 

Simpson, A. A., C.M.G., Lockwood Road, Burnside. 

Snow, FRANCIS (oles, National Mutual Buildings, King 
William Street. 

*“Sranutey, E. R., Government Geologist, Port Moresby, 
Papua. 

Sutton, J., Fullarton Road, Netherby. 

*TIEGS, Oscar Wi Mess University of Adelaide. 

*Torr, W. G., 1 Ve Das M.A., B.C.L., Brighton, South Aus- 
tralia. 

*TurneR, A. JEFFERIS, M.D., F.E.S., Wickham Terrace, 
Brisbane, Queensland. 

*“Veroo, Str JosepH C., M.D. (Lond.), F.R.C.S., North 
Terrace, Adelaide. ; 

*Waite, Epear R., F.L.S., Director South Australian 
Museum. 

*Warp, Leonarp Keitn, B.A., B.E., Government Geologist, 


Adelaide. 
WerpensacH, W. W., A.S.A.S.M., Glencoola, Glen 
Osmond. 
Wuirsreap, Howarp, c/o A. M. Bickford & Sons, Currie 
Street, Adelaide. 
“Ware, Caprarn S. A., C.M.B.0.U., ‘““‘Wetunga,”’ Fulham, 


South Australia. 
*Witton, Proressor J. R., D.Sc., University of Adelaide. 


ASSOCIATE. 


Rozsinson, Mrs. H. R., ‘‘Las Conchas,’’ Largs Bay, South 
Australia. 


| 


667 


APPENDIX. 


FIELD NATURALISTS’ SECTION 


OF THE 


Boval Society of South Australia (Sneorporated). 


THIRTY-NINTH ANNUAL REPORT OF THE 
COMMITTEE 


For THE YEAR ENDED SEPTEMBER 26, 1922. 


The Committee has pleasure in recording a year of pro- 
gress and work well done, activities in many branches of 
science being well maintained. 

Last year’s membership was recorded as 132, and there 
have been 63 new members elected, and several resignations 
and deaths during the twelve months, so that our present 
membership totals 183. We have every reason to be gratified 
at this large increase, which it is hoped will continue. 

Excursions.—On the whole, the excursions have been 
well attended and interesting information has been given 
by the various leaders. The subjects have been as follows :— 
Forestry, Geology and Minerals, Zoology, Shore Life, 
Physiography, Shells, Botany, Nature Study, Pond Life, 
Dredging, Botanic Gardens. 

Lectures.—The Lectures have been of the usual high 
character, and we are much indebted to those who gave them. 
The’ subjects have been as follows:—Aquaria, Artesian 
Waters, Native Camps, Conchology, Mushrooms, Entomology, 
Ooldea, ‘‘Through Australia,’’ Crystals, Recently Introduced 
Weeds, and Plant Curiosities. The attendances have been 
generally good, and quite a large number of visitors have been 
present. During the year an innovation was introduced by 
which lectures dealing with the elementary phases of natural 
science were given. More lectures of this kind are needed, 
and the Committee intends giving attention to this for the next 
programme. 

Exuisits.-—_Numbers of specimens were brought to the 
meetings and were always interesting. It cannot be said that 
this item has been given too much prominence, and it is 
hoped that more members will avail themselves of the invi- 
tation to bring exhibits at every meeting. 


668 


Wixtp FLower SHow, 1921.—A very successful show was 
held on September 23 and 24, and the net proceeds amounted 
to £69. 

‘““Toe Sourm AusTRALian Natura.ist.’’—Our paper has 
been published quarterly and has been the means of main- 
taining interest in our Section. 

VERNACULAR Pxrant Names.—The Sub-committee ap- 
pointed has net met during the year. It is understood that 
the Victorian Field,yNaturalists’ Club is publishing a new 


flora of that State, in which common names will be shown.. 


It may be possible to include popular names in the new Flora 
of South Australia now being prepared. 

FLOWER SHOWS IN OTHER StTaTES.—At our previous 
flower shows we have been fortunate in receiving big consign- 
ments from. other States, and we have reciprocated as far as 
possible. This year parcels of native flowers have been sent 
as follows:—(1) To Melbourne, Victorian F.N. Club’s Exhi- 
bition, June 20. (2) To Sydney, Naturalists’ Society of New 
South Wales Exhibition on September 7 and 8. (3) To 
Broken Hill, Barrier Field Naturalists’ Club Wild Flower 
Show on September 9. We intend sending wild flowers to 
the Queensland Naturalists’ Club, September 30, and Vic- 


torian Field Naturalists’ Club, October 3. It has also been — 


arranged to make an exhibit of wild flowers at the Sweet 
Pea Exhibition, in the Adelaide Town Hall, on September 23, 
and at the Horticultural and Floricultural Society’s Flower 
Show on October 27. Mrs. Page, of Myponga, has been a 
great help in this connection. 

NEWSPAPER Reports.—We are grateful to the daily 
papers for inserting our reports of excursions and lectures, 
and to. The Register, in particular, for its sympathetic atti- 
tude generally towards Nature subjects. 

OxzituaRy.—lIt is our sad duty to record the death of 
several members during the term as follows:—Mr. G. De 
Caux, a young man who was deeply interested in Nature, 
and who had made a special study of orchids, and was the 
first to discover in South Australia the Duck Orchid (Caleana 
major). He was studying for the ministry and gave pro- 
mise of exceptional ability. Mr. Jas. Aitken died recently 
at an advanced age, and was known to our Section for his 
wide knowledge of natural history. Mr. A. M. Drummond 
was a member for a number of years, and through his genial 
personality he was well liked. His interests in natural history 
were of a general character. 


Wma. Ham, Chairman. 


Ernest Isine, Hon. Secretary. 


eee 


669 


THIRTY-THIRD ANNUAL REPORT OF THE NATIVE 
FAUNA AND FLORA PROTECTION COMMITTEE 
For THE YEAR ENDED SEPTEMBER 20, 1922. 


Two meetings were held during the year. 


It is to be regretted that the proposed Trees and Road- 
sides Bill, referred to in last year’s report, was defeated in 
Parliament. 

The Minister of Industry was approached by the Chair- 
man in terms of Mr. Bristow’s request to have a Reserve 
for Kangaroos and Emus in the Flinders Range, but he was 
advised that the Ministry were not prepared to take notice 
of the application without its being consented to by the 
landholders affected. Mr. Bristow was accordingly advised to 
get up a petition by those concerned in order to obtain what 
he desired. 

- A report was ene to the Minister of Industry drawing 
attention to owls being kept in captivity by a dealer in this 
city. Action was taken, and we were advised later that the 
birds had been liberated by the dealer. 


The Minister was also informed that opossums were being 
shot near Urrbrae and Netherby. The police were instructed 
to investigate, but owing to the lapse of time they were unable 
to secure the offender. The Minister, therefore, requests that 
prompt intimation be given to his Department in any future 
breaches of the regulations of the Animal and Bird Pro- 
tection Act. 

Notification was sent that shooting at ducks appeared to 
be taking place on the Thorndon Park Reservoir. Action was 
taken by the Department to have that stopped, and the Water- 
works and Sewers Department, under whose control the 
Reservoir is placed, was notified. 

FLInDERS CuHase.—The situation with regard to this is 
progressing, and matters of improvement are now, it is under- 
stood, before the Board. On August 18 last the Chairman 
delivered a lecture at the Town Hall, at which a collection 
was taken up in aid of this Chase and resulted in a fair sum 
being handed over to the Board. 

Through business requirements, Mr. J. Neil McGilp 
relinquished the office of Hon. Secretary, and a very hearty 
vote of thanks was passed by the members of the Committee 
for his past services. 

J. Surron, Hon. Secretary. 

September 9, 1922. 


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{Generic and specific names printed in italics indicate that the 
forms described are new to science. | 


Abraxas sporocrossa, 285 

Acanthocephala in Australian Birds, 
91, 108 

Acanthochites, 579 

Acanthochiton, 9; A. cornutus, 17; 
costatus, 10; coxi, 18; gabrieli, 10; 
jacundus, 578; mayi, 12;  retro- 
jectus pustulosus, 15; shirleyi, 13; 
stewartiana, 579; sueurii, 578; 
violaceus, 578; v. papillo, 578; 
zelandicus, 579 

Acarina of Australian Birds, 99, 115 

Accipitriformes, Parasites of, 85, 89, 
90, 92, 94, 97, 100, 101, 103 

Acidaha, 205; A. despoliata, 265; 
hypochra, 265; synethes, 266; 
perialurga, 266; tenuipes, 266 

Acrostalagmus cinnabarinus, 177 

Adaluma, 557; A. urumelia, 537 

Adamson, R. 8S, and T. G. B. 
Osborn, Ecology of Ooldea 539 

Aeciduim oleariae, 172 

Aelochroma, 281 

Agathia ochrotypa, 277 

Aizoaceae, 597 

Allelidea brevipennis, 318 

Amarantaceae, 596 

Amphipoda, 34 

Anisodes pulverulenta, 269 

Annual Meeting, 615; Report, 650; 
Balance-sheets, 652 

Anseriformes, Parasites of, 88, 89, 
93, 97, 101 

Anthicus strigosus, 298 

Anthotroche truncata, 605 

Anthozoa (Miocene), 138, 140 

Antimimistis, 233; A: illaudata, 234 

Arctocephalus forsteri, 193 

Ardeiformes, Parasites of, 88, 93, 96, 
100,- 101, 103 

Arthropterus articularis, 310 

Asclepiadaceae, 602 

Ashby, E., Notes on Australian 
Polyplacophora, with Descriptions 
of Three New Species and Two 
New Varieties, 9; Types of Aus- 
tralian Polyplacophora described 
by de Blainville, Lamarck, de 
Rochebrune, and Others, 572 

Asterina Baileyi, 175 


Atriplex leptocarpum acuminatum, 
568 

Aureobasidium vitis album, 174 

Australasian Polyplacophora, 572 

Australian Coleoptera, 509 

Australia, Orchidology of, 148 


Babbagia acroptera deminuta, 568 

Bacterium mori, 179 

Balance-sheets, 652 

Balcoracana Creek, Geology of, 74 

Bassia decurrens, 567; limbata, 567; 
paradoxa latifolia, 567; ventricosa, 
566 

Berry, P. A., Investigation of Essen- 
tial Oil from Eucalyptus cneori- 
folia, 207 

Birds, Examined for Entozoa, 109; 
in which Entozoa have not been 
detected, 109; Parasites of Aus- 
tralian, 85 

Black, J. M., Additions to Flora of 
South Australia, 565 

Blinman and Neighbourhood, Geol- 


ogy of, 53 
Boarmia destinataria, 284; maculata, 
284; panconita, 284; pissinopa, 


284; zascia, 283 
Borraginaceae, 602 
Brachiopoda (Miocene), 138, 140 
‘Brown Coal at Moorlands, 

Deposits, 528 
Bursada flavannulata, 286 


HGS 


Caladenia carnea aurantiaca, 154; 


dilatata, 159; gladiolata, 159; 
pumila, 152 
Calldchiton dentatus, 572; platessa 


fossa, 19 
Calochilus paludosus, 156 
Calotis multicaulis breviradiata, 604 
Campanulaceae, 603 
Caryophyllaceae, 597 
Casbia rhodoptila, 288 
Casuarinaceae, 595 
Casuariiformes, Parasites of, 87, 95 
Cats, Wild on St. Francis Island, 
191 
Celerena, 293; C. griseofusa, 294 
Central Australia, Isopod from, 23 


672 


Cephalothecium roseum, 177 

Cercospora apii, 177 

Cestodes in Australian Birds, 87, 104 

Chaetolopha, 243 

Chalcid Wasp, 
319 

Chapman, F., and D. Mawson, The 
Tertiary Brown-coal Bearing Beds 
of Moorlands, 131 

Charadriiformes, Parasites of, 88, 91, 
93, 96, 99 

Chenopodiaceae, 595 

Chenopodium  carinatum  meluno- 
carpum, 506; - microphyllum 
desertorum, 566 

Chiloglottis Gunnii, 159 

Chilton, C., New Isopod from 
Central Australia belonging to the 
Phreatoicidae, 25; Amphipoda and 
Isopoda of Nuyt’s Archipelago and 
the Investigator Group, 34 

Chiton, 575, 574, 575, 579, 581, 582 

Chitonellus, 577 

Chlamydopsis epipleuralis, 310 

Chloroclystis eurylopha, 238; nigri- 
lineata, 239; phoenochyta, 237; 
poliophrica, 240; pyrsodonta, 238 

Chlorocoma melocrossa, 274; nep- 
tunus, 274; rhodothrix, 273; sym- 
bleta, 273; tachypora, 274 

Chrysochloroma, 276 

Chrysocraspeda cruoraria, 268 

Cintractia hypodytes, 174; 
ficus; 1 

Cladosporium phyllophilum, 177 

Cleland, J. B., Parasites of Aus- 
tralian Birds, 85. Exhibit: Puff- 
ball, 613 

Cleora lacteata, 283 

Clepsiphron, 286; C. calycopis, 287 

Coccinellidae, 303 

Coccyges, Parasites of, 94, 98, 101 

Coleoptera, Australian, 309; 
Nuyt’s Archipelago, 295 

Colletotrichium schizanthi, 177 

Columbiformes, Parasites of, 87, 92, 
95, 101 

Compositae, 603 

Coniothecium chromatosporum, 178; 
scabrum, 178 

Coniothyrium acaciae, 176 

Convolulaceae, 602 

Cooper Creek, Cylindro-conical and 
Cornute Stones found at, 304 

Coraciiformes, Parasites of, 89, 90, 
92, 94, 98, 101 

Cornute Stones from the Darling 
River and Cooper Creek, 304 


Metamorphosis of, 


spini- 


of 


Corysanthes, 158 

Crassulaceae, 597 

Cretheis, 230; C. atrostrigata, 231; 
cymatodes, 231 

Cruciferae, 597 

Crustacea, Proterozoic or Lower Cam- 
brian, 6 

Crypsiphona, 279; C. eremnopis, 279 

Cryptoconchus, 576; C. monticularis, 
579; stewartianus, 579 

Cryptoplax laevis, 577; lamarcki, 
576; larvaeformis, 576; montanoi, 
576; striatus, 577; torresianus, 577 — 

Cucurbitaceae, 603 


-Cupressinoxylon in Australian Ter- 


tiary, 535 

Cylindro-conical Stones from the 
Darling River and Cooper Creek, 
304 

Cymatoplex halcyone, 272 

Cymodoce longicaudata, 37 

Cyneoterpna, 279 

Cyperus exaltatus minor, 565 


Dadoxylon in Australian Tertiary, 
535 

Darling River, Cylindro-conical and 
Cornute Stones found at, 304 

Darluca filum, 176 

Wasysternica, 256; D. erypsiphoena, 
257; pericalles, 256 ° 

Dasyuris melanchlaena, 258 

David, T. W. E., Occurrence of 
Remains of Small Crustacea in the 
Proterozoic or Lower Cambrian 
Rocks of Reynella, 6 

Dendrobium dicuphum, 154 

Dermestidae, 296 

Deto marina, 35 

Diastoma melanioides, 607 

Diploctena pantoea, 254 

Diplodia citricola, 176 

Diptera of Australian Birds, 94, 113 

Dirce, 290; D. aestodora, 291 

Diurus aurea; 157; brevifolia, 148; 
longifolia, 157 


Eccymatoge, 243; E. callizona, 243; 
morphna, 243 

Echinodermata (Miocene), 138, 140 

Ecological Notes on South Australian 
Plants, 583 

Ecology of Ooldea, 539 

Ectroma benefica, 295 

Edquist, A. G., Exhibit: Loranthus, 
614 

Eleale aenea, 317; pulchra, 316; sim- 
plex, 318 


nemiocinosneiiid 


_ Elston, A. H., Australian Coleoptera, 
309 

Entozoa, 116 

Eois albicostata, 261; 
264; chloristis, 263; 
delosticta, 264; elachista, 263; 
epicyrta, 263; ferrilinea, 261; 
miltophrica, 262; prionosticha, 263; 
scaura, 262 

Eremophila pentaptera, 570 

Eriachne ovata pedicellata, 565 

__ Erysiphe cichoracearum, 175 

_ Eucrostes, 272; E. iocentra, 272 

Eucalyptus cneorifolia, 207 

Eucela, 276; E. amalopa, 276 

Kucyclodes, 277; E. dentata, 277 

Euloxia argocnemis, 273; gratiosata, 
272 

Euphorbiaceae, 600 

Kuphyia, 248; E. 
coniophylla, 253; 
oxyodonta, 251; panochra, 251; 
perialla, 249; poliophasma, 252; 
symmolpa, 250; 
tacera, 248; 


argophylla, 
costaria, 261; 


aprepta, 253; 
leptophrica, 250; 


trissocyma, 252 


Field Naturalists’ Section, 667 

Flinders Range, Geology of, 46 - 

Flora and Fauna of Nuyt’s Archi- 
pelago and the Investigator Group, 

_ No. 1, Amphipoda and Isopoda, 34; 
No. 2, Monodelphian’ Mammals, 
181; No. 3, Sketch of the Ecology 

of Franklin Islands, ahh No.. 4, 
Coleoptera, 295 

Flora of South Australia, Ackditions 
to, 565 

Foraminifera (Miocene), 138 

Frankeniaceae, 601 

Franklin Island Rat, 181 

Franklin Islands, Ecology of, 194 

Fumago vagans, 178 


Galliformes, 
100 
Gasteropoda (Miocene), 140 


epimitra, 275; iseres, 275; 

pasta, 275; orthodesma, 276 

Geometrites, 225 

_Geraniaceae, 600 

Gloeosporium ribis, 177 

~Gnamptoloma chlorozonaria, 268 

Goodeniaceae, 603 

_ Gramineae, 594’ 

Grindstone Range, Flinders Ranges, 
74 

Grubia setosa, 35 


lychno- 


oad 


673 


symphona, 248; 


Ischnochiton campbelli, 574; 


Parasites of, 89, 91, 95, 


Gelasma, 274; G. centrophylla, 276;- 


Gruiformes, Parasites of, 93, 96 

Gymnoplax adelaidensis, 582 

Gymnoscelis acidna, 235; holocapna, 
257; Kennti, 236; lophopus, 235; 
spodias, 235; subrufata, 235; 
tanaoptila, 235 

Haematozoa, Australian 
Birds, 100 

Haliplidae, 309 

Halorhynchus caecus, 302 

Harpographium corynelioides, 178 

Helaeus castor, 298; modicus, 298 

Helicella ventricosa, 609 

Helicopage cinerea, 278 

Hemichloreis theata, 279 

Horisme, 244; H. mortuata, 244; 
plagiographa, 244 

Howchin, W., Geological Traverse of 
Flinders Range from Parachilna 
Gorge to Lake Frome Plains, 46. 
Exhibits: Glaciated erratics from 
Central Australia, 612 

Hyperomma lacertinum, 295 


TER;) of 


idicchra, 2i0:. ¥ 
demissa, 271 

Idiodes argillina, 289 

Insect Metamorphosis, 319 

Investigator Group, Flora and 
Fauna of, 34 


celidota, 271; 


lineo- 
latus, 573; longicymba, 573; 
melanterus, 574; sulcatus, 574; 
tessellatus, 574 

Ising, E. H., Ecological Notes on 
South Australian Plants, 583 

Isoodon barrowensis, 39 

Isopod from Central Australia, 23 

Tsopoda, 35 

Iulops, 272 


Jack, R. L., Exhibit: Model of Iron 
Knob and Vicinity, 613 

Janjukian (Miocene), Fossils from, 
137 

Johnson, E. A., Exhibit: Unio, 613 

Jones, F. W., External Characters 
of Pouch Embryos of Marsupials, 
Isoodon barrowensis, 39; Pseudo- 
chirops dahlia, 119; Monodelphian 
Mammals of Nuyt’s Archipelago 
and the Investigator Group, 181. 
Exhibits: Bones of Thylacoleo and 
Thylacinus, 611; Myrmecobius, 613 


Kalimnan (Lower Pliocene), Fossils 
from, 136 


674 


Kellermannia pruni, 176 
Kimber, W. J., Exhibits: 
from Point Turton, 612 

Kochia seleroptera, 568 


Fossils 


Labiatae, 602 
Larentia aganopis, 246; oribates, 
246; petrodes, 245; xerodes, 245 

Lariformes, Parasites of, 93, 96 

Lea, A. M., Coleoptera of Nuyt’s 
Archipelago, 295. Exhibits: In- 
sects, 610, 611, 612, 613, 615; Owl 
pellets, 611 

Lecanomerus flavocinctus, 295 

Leguminosae, 598 

Lemidia alternata, 318 

Lepidopleurus, 574; L. fodiatus, 572 

Lepidoptera, Australian, 225 

Leporillus jonesi, 183 

Leucothoe spinicarpa, 34 

Library, Donations to, 654 

Liliaceae, 595 

Liolophura gaimardi, 581; 
581; hirtosa, 579 

Loboplax, 579 

Loranthaceae, 595 

Loricella angasi, 22 

Lower Cambrian Rocks, 6 

Lycaeninae, New Genus and Species, 
537 

Lycosa perinflata, 84; 


georgiana, 


skeeti, 83 


Mallee, Narrow-leaf, 207 

Mallophaga of Australian Birds, 95, 
113 

Malvaceae, 600 

Mandalotus lutosus, 318 

Marsiliaceae, 594 

Marsupials, Pouch Embryos of, 39, 
119 

Mawson, D., Calcareous 
from Caves, 610 

Mawson, D., and F. Chapman, Ter- 
tiary Brown-coal Bearing Beds of 
Moorlands, 131 

Meadows Valley, 
Features of, 160 

Meetings, Ordinary, 610; 
615 

Melitulias, 247; M. leuwcographa, 247 

Members, List of, 664 

Menuriformes, Parasites of, 92, 95, 
98, 100 

Mesembrioxylon in Australian Ter- 
tiary, 530 

Metallochlora neomela, 277 

Metoponorthus pruinosus, 37 

Micrectyche nana, 298 


deposits 


Physiographical 


Annual, 


Microdes, 240; M. 
oriochares, 240 
Microfilariae in Australian Birds, 90 

Minoa, 233 

Miscellanea, 607 

Mites of Australian Birds, 115 

Mixocera, 272 

Monodelphian Mammals, 181 

Moorlands Brown Coal, 528 

Moraea xerospatha monophylla, 566 

Mount Chambers Creek, Geology of, 
70 : 

Mount Lyail, Geology of, 70 

Myoporaceae, 602 

Myrtaceae, 601 


asystata, 241; 


Nasonia, 319 

Nematodes in Australian Birds, 89, 
107 

(New Zealand, Orchidology of, 148 

‘Nobes, E. D., Preliminary Note on 
Fossil Woods from some Aus- 
tralian Brown Coal Deposits, 528 

‘Noreia loxosticha, 293 

Notoplax, 10 : 

‘Nuyt’s Archipelago, Flora and 
Fauna of, 34, 181, 194, 295 


Obituary, F. R. Zietz, 610 

Obolella Limestone, 67 

Oenochroma artia, 292; lissoscia, 292 

Oidium, 175; O. oxalidis, 178 

Onithochiton astrolabei, 582; lyelli, 
582; neglectus, 582; undulatus, 582 

Ooldea, sEcology of, 539 

Orchidology of Australia and New 
Zealand, 148 

Ordinary Meetings, 610 

Osborn, 2 eGs 'B., Pathological ~ 
Morphology of Cintractia spinificis, 
1; New Records of Fungi for ~ 
South Australia, together with a 
Description of a New Species of 
Puccinia, 166; Ecology of Franklin 
Islands, 194 b 

Osborn, T. G. B., and R. S. Adam- | 
son, Ecology of Ooldea, 539 

Osborn, T. G. B., and G. Samuel, | 
Some New Records of Fungi cle 
South Australia; together with a 
Description of a New Species of 
Puccinia, 166 ty 


2 _ 
Pamphlebia, 274; P. rubrolimbaria, 
274 
Papaveraceae, 597 ; 
Parachilna Gorge, Geology of, 47 ¢ 
Parasites of Australian Birds, 65, 104 
by 9 


@ 


i? Z| 


- Paridotea ungulata, 36 
- Parodiella banksiae, 175 
Passeriformes, Parasites of, 89, 90, 
= 92, 94, 95, 99, 100, 101, 103 
Patawarta Hill, Geology of, 60 
Pelecaniformes, Parasites of, 85, 89, 
90, 94, 97 
Pelecypoda (Lower Pliocene}, 
(Miocene), 138, 140 
Pentarthrocis, 302; P. ammophilus, 


503 


136; 


Perixera flavirubra, 268; lapidaita, 
268 

Phiogistus agraphus, 312; Jleuco- 
cosmus, $14; punctatus, 315; 


rubriventris, 313; ungulatus, 314 

| Phoma macrophoma, 176 

_ Phréatoicidae, 23 . 

Phreatoicus latipes, 26 

Phyllosticta brassicicola, 176 

Phytolaccaceae, 597 

Picrophyila, 287; P. hyleora, 288 

_ Pingasa, 280; P. acutangula, 280; 

| atriscripta, 281; muscosaria, 280 

| Pisces (Lower Pliocene), 136 (Mio- 
cene), 138, 140 

Pisoraca simplex, 269 

Pittosporaceae, 598 

Plantaginaceae, 603 

Plasmopora viticola, 178 

Plaxiphora albida, 575; 
515; costata, 575; glauca, 
varipilosa, 576 

Podicepiformes, Parasites of, 87 

Poecilasthena, 231; P. panapala, 232; 
sthenommata, 252; thalassias, 232; 
xylocyma, 232 

Polygonaceae, 595 

Polyplacophora, 9; Australasian, 572 

Polypodiaceae, 594 

Polyzoa (Miocene), 138, 140 

Porcellio laevis, 37 

Portulacaceae, 597 


biramosa, 
Sia 


Pouch Embryos of Marsupials, 39, 


115 

Prasophyllum australe, 158; a. vis- 
cidum, 154; Brainei, 149; brevi- 
labre, 158; Frenchii Tadgellianum, 
153; Suttonii, 157 

' Primulaceae, 601 

_Procellariiformes, 
89, 96 

- Proteaceae, 595 

) Proteaceous Plants in Tertiary, Mur- 

_ vay Plains, 145 

_ Proterozoic Rocks, 6 

‘Prototypa dryina, 268 


Parasites of, 88, 


hi 
ie 


Pseudochirops dahli, 119 

Pseudomonas juglandis, 179 

iPsittaciformes, Parasites of, 88, 90, 
97, 100, 101 

Pterohelaeus nitidissimus, 298; ovalis, 
298; simplicicollis, 298 

Pterostylis cycnocephala, 158; hum- 


alis, 151; Mlitchelli, 158; pedo- 
glossa, 158; pyramidalis, 158; 
rufa, 158 


Puccinia angustifoliae, 169; bromina, 
168; calendulae, 170; erechtites, 
171; flavescentis, 169; hibbertiae, 
171; operculariae, 171; saccardoi, 
169; semibarbatae, 169;  vitta- 
diniae, 171 

Pulleine, R. H., Two New Species of 
Lycosa from South Australia, 33; 
Cylindre-conical and Cornute Stones 
from Darling River and Cooper 
Creek, 304 

Pyrenochaete rosella, 176 


Rabbits on Flinders Island, 191 

Ralliformes, Parasites of, 93, 96 

Rats on Franklin Island, 181 

Rhyssoplax canaliculatus, 579 

Rogers, R. S., Contributions to 
Orchidology of Australia and New 
Zealand, 148; Presidential Address, 
615 

Rubiaceae, 603 


‘Samuel, G., and T. G. B. Osborn, 
Some New Records of Fungi for 
South Australia; together with a 
Description of a New Species of 
Puccinia, 166 

Santalaceae, 595 

Sapindaceae, 600 

Saragus' brunnipes, 
297; posidonius, 296 

Sauris perophora, 230 

Scarabaeidae, 296 

Scheonus tesquorum, 565 

Scheuchzeriaceae, 594 

Scopodes sigillatus, 295 

Scotocyma, 242; S. albinotata, 242; 
euryochra, 242; idioschema, 242 


298 ; 


oleatus, 


‘Seydmaenus franklinensis, 295 


Sea Lions on Islands of South Aus- 
tralia, 192 

Seals on Islands of South Australia, 
192 

Septoria depressa, 176; dianthi, 176; 
lepidii, 176; lycopersici, 177 

Seynesia banksiae, 175 


Siphonaptera of Australian Birds, 
94, 113 

Solanaceae, 602 

_ Sphenisciformes, Parasites of, 94, 96 

Spiranthes australis, 155 

Stackhousiaceae, 600 

Stenochiton longicymba, 573 

Sterictopsis, 282 

Sterigmatocystis nigra, 178 

Sterrha ewclasta, 267; ooptera, 267 

Stipa eremophila dodrantaria, 565; 
pubescens comosa, 505;  setacea 
latiglumis, 565. 

Strigiformes, Parasites of, 89, 92, 94, 
101, 103 

Symmimetis muscosa, 234; 
254 

Synchytrium taraxaci, 179 

‘Sypharochiton maugeanus, 21; pellis- 
serpentis, 20, 579; sinclairi, 20 


sylvatica, 


Tarsostenus univittatus, 315 
Tarsotenodes leucogramma, 316 


Teale, HE. 0... Physiography ot 
Meadows Valley, Mount Lofty 


Ranges, 160 

Terpna, 281; T. hypochromaria, 282 ; 
unitaria, 281 

Tertiary Brown-coal Bearing Beds of 
Moorlands, 1351 - 

Thelymitra grandiflora, 157; longi- 
folia, 157; ). Macmillan t56; 
megcalyptra, 156; urnalis, 157 

Thymelaeaceae, 601 ~ 

Ticks of Australian Birds, 115 

Tiegs, O. W., Arrangement of Stria- 
tions of Voluntary Muscle Fibres 
in Double Spirals, 222; On the 
Structure and  Post-embryonic 
Development of a Chalcid Wasp, 
Nasonia, 319; On the Physiology 
and Interpretation of the Insect 
Metamorphosis, 319 

Timareta crinita, 299; hamata, 300; 
incistpes, 301 

Tindale, N. B., New Genus and 
Species of Australian Lycaeninae, 
557 


676 


Yallourn Brown Coal Deposits, 532 


Todima fulvicincta, 310 

Turner, A. J., Australian Lona 
tera of the Group Geometrites, 225 

Trematodes in Australian Birds, 92, 
109 my 


Uldinia, 568; U. mercurialis, 569 
Umbelliferae, 601 - | 
Urocystis hypoxidis, 174 = || 
Uromyces bulbinus, 167; danthoniae, — 

166 q 
Uromycladium tepperianum, 168 
Urticaceae, 595 


Ustilago cynodontis, 173: tepperi, 
173 | ™ || 
3 


Verco, J. C., Exhibits: Snails, oa 
614 4 t 
Vermicularia angustispora, 177; ire 


cinans, 177; varians, 177 


Waite, E. R., Exhibit: Model of 
Camarasaurus, 613 

Ward, L. K., Exhibits: Lante 
Slides of Bucla Basin and Nullar- 
- bor Plain, 610 

White, S. A., Exhibits: Botanieal 
specimens, 610, 611; Birds, 615 

Wilkawillina Gorse! on a 


Geology of, 74 
Wirrealpa and 
Geology of, 65 


Nei ghbourhood, 


Woods, Fossil, 528 
Xanthorrhoe epia, 255; metoporina, 4 


255; sodaliata, 254 . 
Recome 289; X. metallica, 2905 
rubra, 290 
Xylodryas, 285; X. leptoxantha, 286 


Zephyrne, 302 

Zietz, F. R., Obituary of, 610 
TFuzara venosa, 37 

Zz eophyllaceae, 600 


rans. and Proc. 


Roy. Soc. S. Austr., 1922. Vol. XIV Ll, Plate I. 


Fig. 2. 


Pigs i. 


Gillingham, Swann & Co. Ltd., Printers, Adelaide. 


| Trans. and Proc. Roy. Soc. S. Austr., 1922. Wok EVA. Plate Tf. 


° 03 02 03 a4 05 08 OF 08 09 | — 


© Of 02 03 04 05 04 07 08 09 10 
<———ONE MiLLIMETER— — ——> 


Gillingham, Swann & Co. Ltd., Printers, Adelaide. 


anid Proc. Roy, Sec. S. Austr.,1922. Vol. XLVL, Plate IIL 


Gillingham, Swann & Co. Ltd., Printers, Adelaide. 


I-SECTION ACR 


BY WALTER Hoy 


Snake Bend, 


TCROPS FROM THE MOU 


16 


ECTION FROM BLINMAN 


4 ci 


H 


SKETCH-SECTION FROM ' 


er ae 


Trans. and Proc. Roy. Soc, S. Austr., 1922. : 
: Vol. XLVI., Plate IV. 


GEOLOGICAL SKETCH-SECTION ACROSS THE FLINDERS RANGES 
BY WALTER HOWCHIN 


W. 


Outer Escarpment 


Archaeocyathinae 
Limestone 


Parachilna 
Plain 


Inner Escarpment 


The Dairy 


Horne’s 
Camp 


> Reservoir Hill 


, Parachilna Creek 


Snake Bend, “The Big Hill” 
(7-mile Hill) 


Fig. 1.—_GEOLOGICAL SKETCH-SECTION OF OUTCROPS FROM THE MOUTH OF THE PARACHILNA GORGE TO THE VICINITY OF BLINMAN, 
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Fig. 2. Area on south coast of Eastern Franklin, showing sand 
drifting away from and exposing travertine pavement. The 
blown sand is held by Nitraria Schoeberi. 


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bseand Proc. Roy. Soc. S. Austr., 1922. Vol XE. Platetx. 


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Fig. 2. Recent blow-out exposing travertine pavement in 
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CONTENTS. 
Ossorn, Pror. T. G. B.: A Note on the Pathological | a 
_ Morphology of Cintractia spinificis (Ludw.). Plate 1. 1 man 
Davin, Pror. Sir T. Engeworts: Occurrence of Remains of a 
Small Crustacea in the Proterozoic(?) or Lower Cam- a 
brian(?) Rocks of Reynella, near Adelaide. Plate il, ... ~~ “gf 
AsHBy, Epwin: Notes on Australian Polyplacophora, with 
Descriptions of New Species. Plate il. ... spp 4" 
Cuitton, Pror. CHaRLES: A New Isopod from Central Aus- 
tralia belonging to the Phreatoicidae _... nino i) Me 
Cuinton, Pror. Cuarues: The Flora and Fauna of Nuyt’s 
Archipelago and the Investigator Group, No. 1—The 
Amphipoda and Isopoda iy ANS. 4 SVN A fs 
Jonzs, Pror. K. Woop: The Kxternal Characters of Pouch 
Embryos of Marsupials, No. 3—Isoodon burrowensis 
Howcuin, Pror. Waiter: A_ Geological Traverse of the — 
Flinders Range from the Parachiina Gorge to the Lake 
Frome Plains. Plate iv. ... oe a oie ee 
Putueine, Dr. R. H.: Two New Species of Lycosa from South © 
Australia. Plate v. ... Pea Meee ik © Peek We 
CLELAND, Pror. J. Burton: The Parasites of Australian Birds’ 
JongEs, Pror. F. Woop: ixternal Characters of Pouch Embryos Hees 
of Marsupials, No. 4—Pseudochirops dahli, Plate vi.  W9 
Mawson, Pror. Sir Dovetas: The Tertiary Brown-coal ie 
Bearing Beds of Moorlands ... a soe ne oe 
Rogers, Dr. R. S.: Contributions to the Orchidology of 
Australia and New Zealand ... ed Be: sc Sots Coe 
Treats, Dr. KE. O.: The Physiography of the Meadows Valley 160 — 
Osporn, Pror. T. G. B., and Grorrrey SamMuEu: New Records 
of Fungi for South Australia, Part II., with a Descrip- 


a2 eB eke 


eae ; 
Sek 


tion of a New Species of Puccinia. Plate vii. ... fie 
Jones, Pror. F. Woop: The Flora and Fauna of Nuyt’s © 
Archipelago and the Investigator Group, No. 2—The 
Monodelphian Mammals Hiss EAS Bis igi ae eee 
Ossorn, Pror. T. G. B.: Flora and Fauna of Nuyts © 
Archipelago, No. 3—A Sketch of the Ecology of Franklin © 
Islands. Plates viii. to xi. ... sa a Bp, wns GES 
Berry, Puuir A.: An Investigation of the Essential Oil (> 
from Eucalyptus cneorifolia, DC... Bote 


~Tizes, O. W.: On the Arrangement of the Striations of ~~  — 
Voluntary Muscle Fibres in Double Spirals. Plate xii. 222 
Turner, Dr. A. JEFrreris: Australian. Lepidoptera of the — 
Group Geometrites ay am Hy “fas a ps 
Lea, A. M:: The Flora and Fauna of Nuyt’s Archipelago and 
the Investigator Group, No. 4—Coleoptera. Plate xiii, 295. 
PULLEINE, Dr. Ropert: Cylindro-conical and Cornute Stones 
from the Darling River and Cooper Creek. Plate xiv. 304 


Euston, Aupert H.: Australian ’Coleoptera, Part III. ... 309” | 
Tizas, O. W.: Researches on the Insect Metamorphosis. 
Plates xiv. to xxix. . 319 


Nozes, E. Dorotuy: A Preliminary Note on the Fossil 
Woods from some Australian Brown Coal Deposits ... 528 
TinpaLtE, Norman B.: On a New Genus and Species of Aus- 
tralian Lycaeninae. Plate xxxi. ... Hi the we. OBE 
Apamson, R. S., and Pror. T. G. B. Ossorn: On the Ecology 
of the Ooldea District. Plates xxxii. to xxxvi. ... oh 
Biack. J. M.: Additions to the Flora of South Australia, 
No. 20. Plate xxxviil. Wd wee ah ap ... 665 
Asusy, Epwin: Types of Australasian Polyplacophora now in 
the Museum d’ Histoire, Naturelle, in Paris eat vei 
Istne, E. H.: Ecological Notes on South Australian Plants, 
Part I. Plates xxxviii. to xlii. ... ee ae «+: OBO 
MIscELLANEA, 607; ApsTRACT oF Procespines, 610; Presi- 
DENTIAL AppRESs, 615; ANNUAL Report, 650; BALANOE- ° : 
SHEETS, 652; Donations To Liprary, 654; List or MeMperRs 664 — 
AprENDIcES—Field Naturalists’ Section: Annual Report, etc. 667 
Thirty-third Annual Report of the Native Fauna and 
: lora Protection Committee  ... iy mh 


4 Jit 


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